Wild Mammals of Wisconsin (Faunistica)

THE WILD MAMMALS OF WISCONSIN Charles A. Long

INTRODUCTION 1 This page intentionally left blank

2 THE WILD MAMMALS OF WISCONSIN The Wild Mammals of Wisconsin

Charles A. Long

SOFIA–MOSCOW 2008

INTRODUCTION 3 THE WILD MAMMALS OF WISCONSIN Charles A. Long Professor Emeritus at the University of Wisconsin – Stevens Point, Stevens Point, Wisconsin

A E N TA IM N A A LI PL A

FUN

PROTISTA

MONERA

Publication No. 56 Museum of Natural History University of Wisconsin-Stevens Point Reports on the Fauna and Flora of Wisconsin* *Also published as Reports of the Museum, and Reports on the Fauna and Flora.

Publication Costs and the Museum’s Distribution of this Book were funded by Harold Roberts – Stevens Point, Wisconsin

First published 2008 ISBN 978-954-642-313-9 (paperback) ISBN 978-954-642-303-0 (hardback) Pensoft Series Faunistica No 68 ISSN 1312-0174

© PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner.

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4 THE WILD MAMMALS OF WISCONSIN CONTENTS

PREFACE 7

ACKNOWLEDGMENTS 9

INTRODUCTION 11

THE WILD MAMMAL FAUNA 11

IMPORTANCE OF MAMMALS 13

PLAN OF THIS BOOK 17

SCOPE AND PLAN 17

FREE EXPRESSION IN TAXONOMY 21

PRESERVATION OF SPECIMEN COLLECTIONS AND HABITATS 21

COUNTIES AND TYPE LOCALITIES 22

EARLY MAMMALOGY AND NOW 25

ENVIRONMENTS AND ECOLOGY OF MAMMALS 31

GENERAL ECOLOGY OF WISCONSIN MAMMALS 31

FORMER GLACIATIONS 32

CLIMATE 33

PHYSIOGRAPHY 34

SOILS 36

COMMUNITIES 38

MICROHABITATS 41

ECOLOGICAL SUCCESSION AND LAND USE 43

ZOOGEOGRAPHY 47

PREHISTORIC MAMMALS 55

REINTRODUCED AND EXTIRPATED MAMMALS OF WISCONSIN WITH HYPOTHETICAL

LIST OF SPECIES POSSIBLY IN WISCONSIN 59

CHECK-LIST OF WISCONSIN’S WILD MAMMALS 64

KEY TO ORDERS AND FAMILIES OF WISCONSIN MAMMALS 66

TAXONOMIC ACCOUNTS OF MAMMALS OF WISCONSIN ARRANGED IN ORDERS,

FAMILIES, GENERA, SPECIES AND RACES 68

OPOSSUM (ORDER MARSUPIALIA) 68

INTRODUCTION 5 INSECTIVORES (ORDER INSECTIVORA) 75

MOLES (TALPIDAE) 76

SHREWS (SORICIDAE) 84

BATS (ORDER CHIROPTERA) 108

HARES AND RABBIT (ORDER LAGOMORPHA) 143

RODENTS (ORDER RODENTIA) 155

WOODCHUCK, CHIPMUNKS, GROUND, RED, GRAY,

FOX AND FLYING SQUIRRELS (SCIURIDAE) 156

NORTH AMERICAN BEAVER (CASTORIDAE) 220

POCKET GOPHER (GEOMYIDAE) 229

MURID MICE AND RATS (MURIDAE) 235

JUMPING MICE (ZAPODIDAE) 304

NORTH AMERICAN PORCUPINE (ERETHIZONTIDAE) 315

CARNIVORES (ORDER CARNIVORA) 321

COYOTE, WOLF, AND FOXES (CANIDAE) 321

BLACK BEAR (URSIDAE) 351

RACCOON (PROCYONIDAE) 359

MARTENS, WEASELS, MINK, SKUNKS, RIVER OTTER,

NORTH AMERICAN BADGER (MUSTELIDAE) 366

CATS (FELIDAE) 427

DEER (ORDER ARTIODACTYLA) 438

WHITETAIL, MOOSE (CERVIDAE) 439

GLOSSARY 453

LITERATURE CITED AND BIBLIOGRAPHY 461

APPENDIX A. Longevities of Some Mammals That Occur in Wisconsin 528

APPENDIX B. English and Metric Measurements 529

APPENDIX C. Preparation of the Mammal Study Skin with Label and Skull 530

APPENDIX D. Dental Records 533

INDEX TO SCIENTIFIC AND VERNACULAR NAMES OF WILD MAMMALS OF

WISCONSIN 539

6 THE WILD MAMMALS OF WISCONSIN PREFACE malogists in Wisconsin, and in many other states and provinces, need an updated source The mammals of Wisconsin were studied by my of information about Wisconsin’s mammals. friend, the late Dr. Hartley H. T. Jackson, who For this group, the taxonomy was updated, published his findings in 1961. In the past 45 and the new collection of mammals at Stevens years the fauna has changed. Some mammals Point’s Museum of Natural History was stud- have vanished, others have established new ied and processed. The information obtained populations here, and the names of the mam- here and from other studies provides a source mals, both scientific and popular, have under- book for mammalogical investigations. gone many changes. In Jackson’s day science Second, there are many nature lovers in was more observational and descriptive than Wisconsin who have curiosity about animals. today. Although superficially they may seem For this group, nature and wildlife provide the similar, his book is different from this one. Jack- joie de vivre. Even many children have a bent son began his work in the early part of this cen- in this direction, and this curiosity and pas- tury. In his later years he carefully studied the sion should be encouraged with source books status of Wisconsin mammals from their rela- about nature. I was one such child myself. My tions with pioneers through the decades of ex- fifth grade teacher, Velma Kane, gave me a ploitation by hunters, and heavy land use by fine book by Frank M. Chapman that set my settlers, farmers and townspeople. His work path. I make no apology for anecdotal and focused on the turn of the century to about 45 even poetic commentary, for those apprecia- years ago. This work reviews Wisconsin mam- tive of that are a welcome audience. Hope- malogy in light of many new scientific findings, fully this book will serve the nature lover in and at the turn of another new century, indeed, pursuit of enjoyment, enthusiastically finding at the beginning of a new millennium. a “joy for life”. In the presentation of natural history, with Third, professional and amateur conser- particular emphasis on the sign of mammals vationists need factual information on the sta- observed in the field, on material of historical tus of Wisconsin wildlife. They must convince and popular nature, and on the clear descrip- politicians and bureaucrats of ways and needs tions of specimens and species, Jackson has to save wildlife in Wisconsin. Human popula- no peer except perhaps Ernest Thompson tion growth and expanding land use have re- Seton. The bibliography in Jackson’s work duced and altered wildlife habitats through- (1961) is a supreme Wisconsin compilation out the State. In my travels on this continent up to that time. However, there is no men- and abroad I have always been impressed with tion of many modern concepts, even echolo- the natural beauty of our own Wisconsin. cation in the bats; there is too great a depen- There is much wonder and aesthetic pleasure dency on temperature zones to explain the here at home, in Wisconsin, but how little of ecological and geographical distribution of it there is to go around. How easy it seems Wisconsin mammals; and there is not a word for people to convert a scenic and natural about the strongest rationale for modern tax- place into an unnatural commercial develop- onomy, including evolution, fossil record, and ment, unsuitable for wild creatures. speciation of taxonomic categories. Neverthe- Some sportsmen serve both as nature less, Jackson’s work is a “tough act to fol- protectors and conservationists. Aldo Le- low.” I will repeat a famous saying, “cosa ben opold was one, and my friend Fran Hamer- fatta e` fatta due volte” [anything worth do- strom was another. Some teachers fit into ing is worth doing it twice]. this category, as do those people who keep This book is written to serve three groups informed on environment while enjoying of readers. First, serious professional mam- nature, apart from aesthetic enjoyment.

PREFACE 7 Hopefully they will in future continue to con- HOW TO USE THIS BOOK serve important habitats for wildlife, because that will be needed. There are special features that help the read- *** er locate particular topics of interest. Anyone curious enough to put forth effort * The Table of Contents at the begin- in nature study will find that nature takes a ning of the book. person into her confidence, and life will be * The Index for species names at the made richer for such effort. This book con- end of the book. tains interesting and useful information on the * A Glossary of mammalogy terms near wild mammals of Wisconsin. I hope that it may the end of the book. stimulate some important study and conserva- * A Check-list of the species accounts tion of this legacy and resource. Though few of the wild mammals of Wisconsin, immedi- people know about it, the scriptures of the Old ately preceding the accounts, which comprise Testament and the Torah provide those of us most of the text. interested with a wise lesson in conservation. * The Plan of the Book explaining the God says we “strangers” must provide “re- format and rationale for the way the explana- demption of the land” (Leviticus 25:24). tion is presented.

8 THE WILD MAMMALS OF WISCONSIN ACKNOWLEDGMENTS any without being unfair. Adrian Wydeven, Tom Jessen and others (Wisconsin Depart- I am sincerely grateful to my family for the sup- ment of Natural Resources) helped early in port they provided for this study. My wife Clau- several ways. Thor Purrinton helped me sur- dine and two sons Alan and John, his son Tyler, vey Detroit Island. Catherine Ladd (Informa- have traveled to all corners of Wisconsin, to all tion Technology) and Teodor Georgiev (Pen- four borders, even into Upper Michigan, Min- soft Publishers) were very helpful. I had the nesota, and Ontario. The family put up with much-appreciated help of two undergraduate me, while I taught heavy course loads and kept students: Nicholee Theiss, and Sarah Miller. at this project for 40 years in Wisconsin. Museum curators allowed me to examine Secondly, I wish to acknowledge the en- specimens or sent them here on loan includ- during, patient, and competent editorial work ing John Paradiso, Robert Fisher, the de- on this book by Professor Lowell L. Getz, ceased J. Knox Jones, Jr., Robert Baker, the Department of Ecology, Ethology, and Evo- deceased E. Raymond Hall, Michael Carle- lution, University of Illinois, Urbana, Illinois. ton, Al Gardner, the deceased Elmer Birney, He was outstanding and necessary. Frank Iwen, Philip Humphrey, Rollin Baker, The Wisconsin Department of Natural Re- Donald Hoffmeister, Scott Craven, the de- sources (Bureau of Endangered Resources) pro- ceased Fred and Fran Hamerstrom, Don E. vided one grant, jointly with the Stevens Point Wilson, and Wendell Johnson. Photographs Bill Cook Chapter, Isaak Walton League and were given by the late Roger Barbour, Matt the U.S. Department of the Interior. This state Lovallo, Frank Iwen, Chuck Pils, Al Cornell, agency and the United States Fish and Wild- D. Shelley, C. Yahnke, Ron Jurawski, R. life Service provided me a grant (1979-1981) Johnson, and Scott McDonald. James Spe- to study Wisconsin bats. The University Per- ro’s (1978) collection of photos, published by sonnel Committee provided several summer Dover Publications, was useful and appreci- grants that paid part of summer salary. It sup- ated. Sylvia Myhre made the drawings of skulls ported my University of Wisconsin Sabbatical, of moles, shrews and bats. If there are help- which I spent at St. Olaf College working on ers overlooked, please feel appreciated. The fractal geometry and chaos theory (in the same U.S. Postal Service permitted the stamps class as my son John). While there, I wrote up showing mastodon and mammoth, both pre- about l5 species accounts and made over l00 historic elephants, 1996. maps for this project. Paul Humke, Math De- Reviewers of various accounts include partment, helped greatly. Faculty who helped Getz, who read them all, Scott Craven, Rob- in various ways include deceased George Beck- ert Freckmann, Jerry Choate, [the late] Karl er (I used his base map for distributions), Philip Koopman, Wayne H. Davis, who both read Bjork, Virgil Thiesfeld, Shelley Jansky, Robert the bats, Stephen Taft, Mark Boyce, Jerry Bell, Frank Bowers, David Hillier, born for com- Wolff, John Whitaker, Jr., Tim Lawlor, Bruce puters, Robert Freckmann, Stephen Taft, Vin- Kohn, W. E. Berg, K. McCaffery, Joe Mer- cent Heig, Sol Sepsenwol, gifted and kind, ritt, Adrian Wydeven, Tom Howard, L. R. Daniel O.Trainer, Mark Boyce, Stan Szczyt- Petersen, and Rebecca Sealfon. I relied on ko, Christopher Yahnke, Isabelle Girard, and the wolf book by Dick Thiel and a paleonto- at the Madison campus Frank Iwen and Scott logical contribution by R. M. West and J. E. Craven. I am grateful to former teachers from Dallman. Hartley H. T. Jackson, deceased, Kansas, especially the late E. Raymond Hall, helped in the beginning years. Robert Wilson, and Horace Hays. Wanting of some quality illustration, I Many students helped making up speci- gathered some drawings from early and his- mens and helping with fieldwork. I cannot list toric sources. Becoming quite interested in

ACKNOWLEDGEMENTS 9 iam T. Hornaday, E. Raymond Hall, W. H. Burt (artist Richard Philip Grossenheider), John Dean Caton, Cantlon and Garry A. Heidt, Baron Georges Cuvier, John Litvaitis, Joseph Holder and Sir John Richardson et alia, James Bee and Hall, Susan E. Smith; Tom Swearingen, Thornton Burgess, Karl Schmidt (artist Walter A. Weber) and artist G. Mutzel in Wood’s Nat. Hist (a favorite book from the author’s Grandmother Long (J. G. Wood, 1865). My son Alan made the muskrat years ago (1985). Virginia Peterson and the Houghton Mifflin Co. gave permission to photocopy two drawings by Roger Tory Peter- Raccoon. Artist Roger Tory Peterson. Courtesy Virginia son. Elizabeth Schwartz and the University of Peterson, her “first choice” for this book. R. T. Peterson and Missouri Press gave permission to use two James Fisher 1955, © RTP and J. M. M. Fisher, reprinted drawings by Charles Schwartz. Other artists by permission of Houghton Mifflin Co. Rights reserved. are credited in the legends herein. While living, my former teacher E. Raymond Hall gave the art, I expanded the search. Hoping for a permission to use several drawings. The styles kind of legacy of former mammalogists, not include stylized scratchboard-sketches, steel only of quoted observations, I resurrected cuts, woodcuts, and zinc engravings. Color some of their art, for old published works are prints were provided by Virgil Beck and Clark becoming rare. Although the artists them- Bronson. selves are not always mentioned, they may This book could not have been published be credited in the works. Included are black without Harold (“Hal”) Roberts. I told him and white works from Anna B. Comstock, more than once that he was “one of the nic- Anna Stryke, and A. MacKinnon, H. H. T. est men that my wife and I ever knew”. Unas- Jackson (artist B.O.M[cMaugh]), Spencer Ful- suming, kind, a true naturalist, he was and is lerton Baird, Charles Cory (artist Leon L. always helpful. Military service interrupted Pray), Elliot Coues, C. Hart Merriam, Gerrit several years of his graduate work at UW- Miller, Jr. (artist Tertzli), Dan Metz, Ernest Madison, where he took every possible course Thompson Seton ( = Ernest E. Thompson), he could from Aldo Leopold and had obtained Vernon Bailey, Elva Hamersrtom-Paulson, D. a masters degree. G. Elliot, W. H. Flower and R. Lydekker, H. All together we hope to express tribute E. Anthony, William J. Hamilton, Jr., Will- to nature, and wildlife poetry and art!

10 THE WILD MAMMALS OF WISCONSIN INTRODUCTION derived “ true mammals” almost lost the older, reptilian quadrate-articular hinge, but not THE WILD MAMMAL FAUNA quite. A consequential attachment of the old, non-functional hinge (both bones already con- “Wild” mammals try to survive in nature with- nected together) to the nearby reptilian hear- out human care, not as provided for “domes- ing bone called the stapes formed a new tic” mammals. They are seldom seen, for most “chain of ossicles.” That is how the derived are secretive and nocturnal. Many spend the “true mammals” came to possess three ossi- day, some the entire winter in burrows. Mam- cles in each ear instead of one. Gould (1990) mals have attractive ellipsoid bodies, with cogently called this story “an earful of jaw”. graceful legs and movements providing swift All three bones, “malleus” (descended from locomotion. Their warm-blooded bodies are the articular), “incus” (from the quadrate) and generally covered with hair (i.e., fur) often rich- even the “stapes” (a transformed hyomandib- ly colored. It may be coarse, felt-like, or soft, ular jaw bone of fishes) were originally jaw and often is marked in tasteful patterns. The bones in ancient fishes. eyes are large, bright, and expressive, except Archaic mammals from the late Triassic in the bats, shrews, and moles. Mammals are Period were probably warm-blooded. Tactile endeared to us because of their interesting bristles, such as those on the scaly tail of the ways and intelligence, to some extent having recent-day opossum, spread over the body behavior almost human. conserving internal heat (Long, 1972b). There Technically, a mammal is a vertebrate an- are two, perhaps more, comprehensive ac- imal noted for its fur (technically called pel- counts on mammalian hair (Ryder, 1973; age) and for the maternal behavior of nurs- Noback, 1951). ing (i.e., suckling) its young. Mammals evolved Long (1972) proposed that Darwin’s spec- long ago from mammal-like reptiles, in the ulation on the origin of lactation in primitive Triassic geological period. Although mamma- mammals was correct. Darwin had believed the ry glands and hair were never fossilized, the mammae first appeared in the pouch. Long fossil record of skeletal transition from reptile believed glandular moistening of the egg and to mammal, especially in South Africa, can young in a pouch provided an ever-improv- be traced out stage by succeeding stage. ing nutrition that became “milk”. This sequence Searching in stratified rocks down into the of associated microevolutionary preadaptations earth, back into the stony records of many fits well with neo-Darwinism. Long (1969c) re- millions of years ago, transitional specimens viewed the evolution of mammary glands. have been found showing a regular trend of Other characters seen only in mammals replacing the archaic, reptilian jaw connec- include a diaphragm to enhance breathing, tion with a brand new jaw joint on each side. a large cerebrum related to intelligence, and Replacement began (of course, on either the loss of the nucleus from every red blood side) by doubling the jaw hinges, i.e., adding cell leaving only a membrane and net-like cy- an articulation between a newly evolved squa- toskeleton of biconcave form containing en- mosal bone of the cranium with the adjacent doplasma and hemoglobin. It has a great sur- dentary bone of the lower jaw. By subsequent- face area for water and gas exchange. This ly reducing the older (quadrate-articular hinge) deformable erythrocyte is driven in blood plas- of each newly doubled jaw joint, while gradu- ma into tiny capillaries, and easily unloads ally enlarging the newer squamosal-dentary oxygen to the tissues (Long,2006). joint, the “new” and “old” joints, paired to- There are 69 to 75 kinds of mammals gether on each side, define those fossils (hav- that are considered wild Wisconsin species. ing “two articulations”) as “mammal-like.” The A species is a population of organisms that

INTRODUCTION 11 breed among themselves, but seldom inter- lease (female or pair) in the Blue River area breed with other species. If hybrids occasion- of two pets brought here from Florida about ally are found, especially those resulting from 25 years ago. pairing in captivity, the two stocks are usually At least three of our mapped Wiscon- regarded as separate species, not necessarily sin species may have recently been extir- as “good” species. The short-tailed prairie pated. Therefore, our recognized fauna (68- deer mouse (Peromyscus maniculatus bair- 69 species) may have declined. Many Wis- dii) possibly in one or two places far away, in consin species have two or even three geo- western states, shows evidence of intergrada- graphic races, called subspecies, so that the tion with long-tailed deer mice. That is actu- number of named kinds is about 79 (a few ally little evidence to prove that this little less than recognized by Jackson, 1961). If mouse hybridizes freely with any race of man- the problematical elk, bison, wolverine, iculatus (which is a long-tailed climbing spe- mountain lion, and caribou, and the minor cies). In any case, bairdii acts as a “good” prairie vole, were all added to the total, the species in the Wisconsin fauna, not hybridiz- number of species rises to 75. The feral ing naturally with long-tailed deer mice, and, house cat, and escaped fallow and Sika deer therefore, is added to the Wisconsin species add species, but are not included; without count. Jackson (1961) did the same; and Bak- human support they would likely disappear, er (1983) described them separately. Some and the two released deer probably have. workers consider the prairie vole race Micro- The house cat is usually a domestic pet. The tus ochrogaster minor to be a valid species, domestic horse (Equus caballus) is not wild and it may be (see account of M. ochrogaster). in Wisconsin. Humans and domestic dogs The introduced European stone marten have long been on the scene in Wisconsin, Martes foina is now established in southeast but are not a wild resource. Wisconsin (Long, 1995). The introduced Nor- The 69 species treated herein, includ- way rat and house mouse were established ing the probable prairie deer mouse spe- long ago. cies, provisional stone marten, and three Removed from this Wisconsin faunal list possibly extirpated species (not found in re- are the native elk Cervus elaphus [C. e. ca- cent years), are classified into distinct ac- nadensis Erxleben], bison Bison bison, wol- counts. Some of these species have no close verine Gulo gulo [= G. g. luscus (Linnaeus)], relatives; others are grouped in “taxa” of mountain lion Felis concolor [F. c. schorgeri two or more species of similar form. These Jackson], and caribou Rangifer tarandus [R. groups are called genera (the singular is ge- t. caribou (Gmelin)] because they were erad- nus). There are about 49 genera; some rec- icated at the turn of the 20th century. The elk ognize Microsorex as distinct from Sorex, was reintroduced (see Reintroductions) more Eutamias as distinct from Tamias, and Pit- than once, and now is protected on a pre- ymys as distinct from Microtus. There is serve in northwest Wisconsin, as well as many controversy whether or not to call the Can- hunting preserves. The bison is now raised as ada lynx and bobcat Felis instead of Lynx, a domestic beef animal on many game farms either use does not change the number of in Wisconsin. The mountain lion, also called genera. Neovison for mink adds one. cougar (Felis concolor), probably has been The genera are classed in 17 families be- released by unauthorized individuals leading longing to 7 orders. These orders are the prim- to some of the numerous reports of cougars itive Marsupialia and Insectivora, aerial bats in Wisconsin; however, there is no verified Chiroptera, rabbit and hares Lagomorpha, population, or even a recent specimen or abundant rodents Rodentia, famous meat-eat- photo of the cougar. I learned of a likely re- ers Carnivora, and hoofed Artiodactyla.

12 THE WILD MAMMALS OF WISCONSIN IMPORTANCE OF MAMMALS Some mammals carry deadly diseases (Ta- ble Intro-1; Davis et al., 1987), which they Mammals are important to the people in Wis- transmit directly or indirectly by parasitic ticks consin. On the positive side, they help fulfill and fleas (such as plague, Rocky Mountain human needs for nature. They provide aes- spotted fever, and Lyme disease). Rabies in thetic experiences, i.e., life-long memories Wisconsin is caused by a virus known in dogs, fondly recalled. They have provided substance cats, cattle, foxes, bats and especially striped for literary anecdotes, legends and tales in- skunks (Table Intro-2). Rabies, also discussed spiring intellectual communication among below under bats (Chiroptera), foxes, rac- people. They furnish highly valued recreation coon, and striped skunk, is a terrible disease for hunters, and flesh to eat. The deer hunt- and always fatal in humans. Usually transmit- ing alone adds millions of dollars to the state ted by mammal bites, rabies can be prevent- economy. Furbearers provide fur for warmth ed by vaccinations. Hantavirus killed one stu- and decoration. Many small mammals serve dent in Kansas, that collected mice. In the as foods for carnivores and raptors, and their years 1996-1997, recent cases include one burrows may be taken over for shelter by other person killed in northern Illinois, and another mammals. The burrows of some mammals person almost killed (April-May, 1997) from (e.g., North American badgers, ground squir- Chippewa Falls, who cleaned a mouse nest rels, pocket gophers, moles, and innumera- out of an old car. Additionally, about 20 kinds ble mice) turn up soil, aerate it, provide nutri- of mammals carry hemorrhagic and arbovi- ents, and facilitate natural soil formation. The rus diseases (Davis et al., 1987), and others beaver turns riparian forestland into vast are involved with plague, tularemia, typhus, meadowlands. There are many mammals con- spotted fever and other diseases (Childs et al., sidered beneficial because they eat insects (as 1995; Mills et al., 1995; Gage et al., 1995; do shrews, moles, especially bats, and to lesser Schmid, 1985). Even wild animals sold as pets extent numerous other species that occasion- may harbor deadly diseases, such as “mon- ally eat them). Some mammals are beneficial key pox” in African rodents and American because they prey on the injurious mice and prairie dogs. rabbits. Much carrion is consumed by opos- Lyme disease is caused by a bacterium sums, carnivores, and many other mammals. Borrelia carried by the ticks Ixodes dammi- On the negative side, rodents and rabbits ni and I. scapularis, and perhaps in other steal our crops and girdle small trees and bush- Ixodes. Untreated, the disease is dangerous. es. The white-tailed deer, lovely and graceful Borrelia is reported in Wisconsin in foxes and as it is, causes over a million dollars damage coyotes (Kazmierczak and Burgess, 1989), to Wisconsin crops annually, hundreds of bears (Kazmierczak et al. 1998), and small thousands of dollars damage to cars, and in mammals (Godsey et al. 1987; Walker et al., recent years unknown but immense costs for 1996). See status in Caporale et al. (2005). damage in urban landscaping. In recent years Arboviruses, those transmitted by arthro- outbreaks of urban deer have devastated gar- pods, include in Wisconsin the La Crosse en- dens and shrubbery. The scriptures call for a cephalitis virus, and probably the St. Louis tithe of our land to nature, but deer, rabbits virus, transmitted by mosquitoes. The La and rodents often eat more than a tithe, some- Crosse virus is reported in red and gray fox- times everything we grow. Although general- es, raccoons, and opossums (Amundson and ly beneficial, weasels, raccoons, skunks, opos- Yuill, 1981 b). Tamias striatus also carries sums, and foxes occasionally may eat the this virus (Gauld et al., 1975), as do Sciurus farmer’s chickens. Some large canids have and Glaucomys (Seymour and Yuill, 1981). preyed on domestic animals. Herpes was reported in cottontails (Schmidt

INTRODUCTION 13 Table Intro-1Intro-1. Important infectious diseases of Wisconsin’s wild mammals. Most can afflict humankind. Additional diseases include tuberculosis, herpes, W. Nile virus, and possibly terrorist dangers such as hoof and mouth and anthrax particularly in our artiodactyls. Rodents are susceptible to plague. Davis et al., 1987, and sources cited in text.

Mammals Rabies Distem- Hemorrhagic Several Tulare- Rocky Brucello- Histo- Other Diseases = X per Diseases Arbo- mia Mtn. sis plasmosis Pseudo- X, Hanta viruses Spotted TB Virus Fever Opossum X X X X St. Louis enc. Some bats X E. cottontail P-TB X X X Lepus P-TB X X Marmota XX Tamias X Eutamias XX Sciurus XX Tamiasciurus X Harvest mouse X Peromyscus X, Hanta virus X X Lyme Disease Clethrionomys P-TB Microtus P-TB X X Ondatra P-TB X Errington’s enc. Porcupine X X Beaver P-TB X X Wolf X X ?X ? Mange, Parvo-virus Coyote, Foxes XX, P-TB XX ?X?X XX ? Mange,Parv. Blastomycosis Raccoon X, P-TB X X X ? Parvo, St. Louis enc. Mink P-TB X X X Lyme dis. tick Weasels X X Mephitis X X X Canine hep. Lontra P-TB Taxidea X X X Plague Martes Martes X Bobcat X X X X Cervus elaphus P-TB X X Odocoileus P-TB X X X X X(rare) Salmonella, Lyme Disease Alces XX X Bison XX Rattus X X X Plague Mus musculis XX XX Felis catus X X X X Mange, Parv.

et al., 1992) and considered in relation to flea Tularemia is an important zoonotic dis- parasitism by Spieker and Yuill (1976). Arbo- ease found in cottontails, hares, beavers, viruses in deer populations were surveyed by muskrats, voles, red fox, gray fox, Muste- Murphy (1989). The recent invasion into the la, and North American badger (Addison et United States, and into Wisconsin, of the West al., 1987). The carnivores often get the dis- Nile virus, carried by mosquitoes, mostly af- ease by scavenging carcasses. Distemper fects birds such as crows. However, it is trans- was found in wild and domestic dogs, the mitted to mammals (e.g, horses, humans) and raccoon, mink, striped skunk, badger, least can cause and already has caused some hu- weasel, and Microtus pennsylvanicus. man mortality. Ringworm was found in juvenile muskrats.

14 THE WILD MAMMALS OF WISCONSIN Table Intro-2Intro-2. Rabies in Wisconsin mammals (1952- 1974). Vaccinations have practically eliminated dog rabies. Bats are low.

Year/bats Skunks Dogs Total Skunk, Bat/Total 1974/ 2 68 17 107 .64//.02 1973/ 9 76 35* 154 .49//.06 1972/ 8 96* 24 163 .59//.05 1971/10* 46 15 91 .51//.11 1970/ 4 33 11 60 .55//.07 1969/ 4 21 6 46 .46//.09 1968/ 6 21 13 54 .39//.11 1967/ 1 28 11 61 .46//.02 Totals 51 389 132 736 .53//.07 1966/ 1 22 — 68 .32//.01 1965/ 1 21 — 64 .33//.02 1964/ 6 12 — 95 .13//.06 Cases of rabies in the United States in wild mammals, 1963/ 5 17 — 62 .27//.08 1955-1991. Krebs et al., 1992. Raccoon incidence is highest in 1962/ 5 12 — 42 .29//.12 eastern states. 1961/ 2 5 — 30 .17//.07 1960/ 0 5 — 24 .21//.00 1959/ 1 64 — 92 .70*//.01 1958/ 3 184* — 227* .81*//.01 1957/ 6 37 — 74 .50//.08 1956/ 0 13 — 41 .32//.00 1955/ 0 19 — 39 .49//.00 1954/ 0 36 — 90 .40//.00 1953/ 0 25 — 49 .51//.00 1952/ 0 27 — 56 .48//.00 Totals 30 499 — 1,053 .47//.03

Anthrax outbreaks, usually arising from soil, Rabies in domestic and wild mammals in Wisconsin are dangerous to many grazing mammals, 1953-1992. Rare since then in domestic animals. and especially those that wallow in mud, such as bison. Unknown in Wisconsin, it is, nevertheless, an imminent threat to man and wolf, at least in some states. The incidence beast, harbored by terrorists and other was reviewed for several species by Short beasts, and is often disseminated by carri- (1989). Vaccination helps control parvovi- on eaters, ranging from gulls to coyotes. rus, protecting our domestic dogs, and has Much the same can be said for bovine TB practically rid the country of rabies in dogs. (= brucellosus) (see Davis et al., 1981). Par- Rapid action by government and the De- voviruses are often lethal to carnivores, es- partment of Natural Resources hopefully can pecially to dogs; this dog virus may have prevent a great calamity following discovery mutated from the feline parvovirus. The ca- (winter 2001-2002) of Chronic Wasting Dis- nine strain of virus went around the world ease in deer near Mt. Horeb. This disease, in the brief interval 1976-1980, providing caused by virus-like “prions” that attack the an example of the incredible cosmopolitan central nervous system, is lethal, and, trans- danger of mutated viruses dispersing in short mitted by unknown means, threatens to dec- times, and the importance of mammals in imate the entire Wisconsin deer population. transmitting viruses. Today it is found in coy- That would create an economic disaster, be- otes, red foxes, raccoons, and probably the cause of the economic importance of deer

INTRODUCTION 15 hunting. See account below of the white-tailed omy and aesthetics, i.e., harmony and beau- deer and see Manwell (2001). ty. If a creature lives in association with hu- The introduced Rattus and Mus defile our mankind, it should be studied carefully by homes and belongings, as well as bringing us people with training so that by scientific man- diseases. The house cat, either feral or do- agement the negatives can be minimized and mestic, eats millions of mammals and our positives can be emphasized. The status and wonderful songbirds (Coleman, 1993). conservation of wild mammals must be con- Any time a mammal reproduces to excess, tinually studied. Then the problems of co-ex- a problem develops, whether the mammal is istence of human and animal may be recog- as small as house mice or as large as the white- nized and solved. Modern-day land use has tailed deer. Usually mammal populations are cost us some irreplaceable species. Wise man- kept in line, especially by predation, diseases, agement has saved some valuable species and and starvation. Raptors certainly do their work replaced others. Today there are famous so- limiting population outbreaks of small rodents. cieties and agencies that set aside habitats, Therefore, biodiversity in nature, a desired goal regulate some populations, and conduct re- of modern conservation, actually benefits the search on mammals that need support. There entire ecosystem and most of its members be- are other agencies, landowners, and users that cause of the so-called “balances” in nature. care little about preservation. They must be Mortality in wild mammals is generally opposed. We should let no mammal, no mat- equal to natality (birth rate). In time they bal- ter how little or secretive it is, escape our ance, so long as populations are stable. Both notice and protection. Those who undertake subjects (mortality and reproduction) are treat- the study of mammals, or any wild creatures ed herein. In nature, potential longevity (see for that matter, will find them far more inter- Appendix A) is usually shortened by natural esting and valuable than they ever might have mortality or hunting and trapping harvests. imagined. Hopefully this work will bring the Most mammals should not be judged as subject of Wisconsin mammalogy, especially bad or good, but managed for a useful func- biology and known status of the wild mam- tion in the community. “Useful” includes econ- mals, up to date, at least to 2007.

16 THE WILD MAMMALS OF WISCONSIN PLAN OF THIS BOOK by, or at some outlier used as a den. The young beaver walks or swims away, the bat flies off, “In pursuing our researches we are often compelled the mole burrows, and the leveret hare jumps to differ from the views of previous writers. . . . we across the prairie. They search for food sourc- will endeavor to be swayed simply by a love of truth, es and den sites, adventures shape their desti- treating all with respect, and adopting such nies, the winds carry the bats to strange plac- language as can be offensive to none.” – John es, and eventually each mammal has sexual James Audubon and John Bachman, The quadru- yearnings. The dispersing young mammal must peds of North America 1842-1846. find a home or resting place, for security, sleeping, and for its offspring. Perhaps it selects a tree cavity, or digs or appropriates a burrow. SCOPE AND PLAN Eventually the mammal may breed and bring forth young hopefully with the same adapta- In this book, the 20th century mammals of tions that served the parents. The genetic pro- Wisconsin are emphasized, but some infor- gram for form and function passes from par- mation is included on prehistoric mammals ents to offspring. and those mammals extirpated during the Thus, it is important for us to determine 19th century. The Prehistoric Mammals are what kind a mammal is (description and discussed in a separate section. The Wiscon- names are provided herein) and where it can sin mammals documented in historic and sci- be found (geographic range or distribution and entific literature but extirpated before 1900, status). The habitat and home are described. and some species that may be found in Wis- Foods, mortality, and reproduction are re- consin but are unverified so far (Hypothetical viewed. Population parameters are discussed, Species, Reintroduced and Extirpated Mam- such as the extent of a species home range mals beyond) are briefly discussed. The treat- and how abundant it is within an area (densi- ment of the mammals is taxonomic; the clas- ty). Under Remarks various subjects are dis- sification is based on evolutionary relation- cussed. Speciation is ascertained for Geo- ships including even the integral concepts graphic Variation. The specimens, mostly in from ecology and physiology. museum collections, are ascribed to county. Unlike humans, fettered but diversified by Some general and characteristic adapta- customs, fads and television, the wild mammal tions, and their phylogeny, are usually dis- lives an odyssey more genetically programmed, cussed for the higher taxonomic categories, common to its kind. Success for a wild mam- such as wings in bats, bunodont molars in mal is survival and successful reproduction. bears, or slenderness in weasels. All of this is Naturalists say the mammal “prefers” a partic- modern taxonomy, and all is based on pale- ular habitat. It occurs in this habitat partly by ontology, physiology, morphology, and ecol- preference. Usually it was born there, of par- ogy, important and inseparable concepts. ents and ancestors adapted to it, as beavers The mammals of Upper Michigan can- are born near water, prairie mammals in grass- not be separated from the Wisconsin fauna lands, or forest mammals amid trees. Since except by the political boundary drawn be- “habitat” means home in a general way, the tween the states. There is no apology neces- den site and nest for most mammals is its im- sary for confining an investigation to an un- portant home within a home. The beginning naturally bounded region, such as a state, but of the odyssey, called “dispersal” by scientists, in my opinion the Upper Peninsula is so con- is departure from the natal den. Actually it is tinuous with Wisconsin ecologically that this weaning, the departure from mother. This re- entire natural region should be appraised as jection may happen in the natal den or near- one. There is continuity of southern Wiscon-

PLAN OF THIS BOOK 17 sin and northern Illinois, and on occasion (e.g., fragment of a population may undergo speci- for Cryptotis, Nycticeius, Pitymys) I also ig- ation leading to the evolution of a new kind. nore that political boundary. Nevertheless, the It follows that some races are differentiated specimens studied herein, in the main, are as quite distinct, as are the Florida and Wis- from Wisconsin, and the book is essentially consin fox squirrels. The prairie and forest about Wisconsin mammals. The east (Lake deer mice in Wisconsin are distinct. Other Michigan) and west (Mississippi River) bound- races seem less distinctive and may be recog- aries of Wisconsin are barriers sharply limit- nized only by careful comparisons of average ing geographic range of some mammals. and significant differences from place to place. Modern comprehensive references for the If I perceive the described geographical dif- surrounding states are Baker (1983) for Mich- ferences, the formal names seem appropri- igan, Hoffmeister (1989) for Illinois, Bowles ate to me. If a former student of geographic (1975) for Iowa, and Hazard (1982) for Min- variation drew a line arbitrarily between two nesota. Wisconsin mammals have been won- different kinds, I may follow his lead. derfully studied in the past by Cory (1912), This book attempts to provide rationales and the late Hartley H.T. Jackson (1961), a for using current scientific (Latinized, post- Wisconsinite formerly of the U.S. Biological Linnaean) names. It broadens the statewide Survey. The reader is urged to consult these study enough to ascertain geographic varia- fine books. No comprehensive work in Wis- tion extending into some other states. There consin has appeared since Jackson’s book, is no aim toward revising the taxonomy of written over 40 years ago. mammal species and subspecies in adjacent The taxonomic sequence of species used states. I searched for resemblance in charac- follows that of Hall (1981), with exceptions ters in neighboring states to assign some Wis- explained in the accounts. Comments describe consin mammals to their taxa. variation on geographic variation, because it The foregoing rationales for formal classi- is such a nebulous concept, and is usually re- fication of geographic variation would not be ferred to subspecies, i.e.races. They appear necessary to convince the scholars Hoffmeis- alphabetically, unless otherwise noted. How- ter (1989) and Jackson (1961), who studious- ever, the nominate race is mentioned first if ly attempted to make careful taxonomic as- present in the state. signments in this region. This raised a peculiar A controversy today in mammalogy is what problem for Wisconsin and Illinois. The polit- to do with geographic races, formally called ical boundary between Illinois and Wisconsin subspecies. These kinds are named and de- unrealistically separates some geographic rac- scribed with trinomens, and too many were es. This dilemma arose in part from inadequa- based on trivial characters. However, some are cies of earlier studies and in part from numer- quite distinctive, and others were discovered ous taxonomists assigning specimens on geo- by carefully appraised comparisons made on graphic grounds to either the nominate sub- specimens collected over a wide region. Once species or the nearest geographic race. Re- considered as incipient species the subspecies spect for colleagues and their use of names at best is a recognizable population more or often resulted in illogical acceptance of a name less adapted to a geographic region. Changes based on trivial or invalid characters. The di- in habitats and other ecological factors may vergent views of the aforementioned two men have affected the gene frequencies and phe- often obtained different results. notypes (appearance) of this struggling popu- Fourteen pairs of subspecies are divided lation, but a distinctive geographic race has by the Wisconsin-Illinois border, called here- genetic properties passed on for generations, in the Hoffmeister-Jackson Phenomena (Ta- usually for thousands of years. An isolated ble Plan-1). They defy the tenets of specia-

18 THE WILD MAMMALS OF WISCONSIN tion (e.g., geographic barriers, or former cli- synonyms and homonyms with their authors’ mates (Long, 1965) that separate evolving names. The citations are sufficient enough, populations). Some reasons for this pattern so they need not be duplicated in the Litera- include (1) natural hiatuses barring population ture Cited (unless they need to be cited else- movements, i.e., valid taxonomy, (2) provi- where in text). The use of parentheses around sional names, and (3) utter speculations. Wis- an author’s name in the synonymy means that consin taxa are listed first. the author named the mammal with a differ- Synonymies are used to head up the spe- ent generic name in the binomen or trinomen cies and subspecies accounts. These list the combination than is presently used. A schol- original use of name for a described kind of ar, then, would never look for the scientific mammal (the scientific name is immediately name Cryptotis parva (Say) in old literature. followed by the Author’s name, which is not In fact, the original combination was Sorex printed in italics). The synonymy also includes parvus Say. changed names, the most modern or recent Most geographic distributions are dis- name combination with the name of the tax- cussed with reference to state maps. The en- onomist who first published that combination. tire North American geographic range of each Obviously all published usages of popular species (after Hall, 1981). is presented in an mammal names today need not to be cited. accompanying map of North America. Where Between the name of the author of a new there is information available, such as for some name combination (not for the original name Wisconsin carnivores, maps based on Depart- itself) and that combination is a colon, because ment of Natural Resources work showing pri- the International Code on Zoological Nomen- mary range, or denoting rare, less common clature recommends that kind of punctuation and common occurrence are often included. be used. To follow that recommendation de- The dots on the state maps are black for spec- fies some precedence in mammalogy. Other imens I have examined, and bull’s-eyes or names listed in the synonymy include invalid open circles for other records from Jackson

Table Plan-1. The Wisconsin-Illinois border separates 14 pairs of subspecies.

Disjunct pairs caused by a natural hiatus, due probably to former climate Synaptomys cooperi cooperi Baird gossii (Coues) Geomys bursarius wisconsinensis Jackson illinoensis Komarke & Spencer* Lontra canadensis canadensis (Schreber) lataxina Cuvier

Races not carefully identified, or given hypothetical names Mus musculus M. m. rutty ? Castor canadensis michiganensis Bailey carolinensis Rhoades Canis lupus lycaon ? Felis concolor schorgeri ?

Problematic taxonomy Sciurus carolinensis hypophaeus Merriam pennsylvanicus Ord Tamiasciurus hudsonicus minnesota Allen loquax (Bangs) Peromyscus leucopus noveboracensis (Fisher) leucopus (Rafinesque) Urocyon cinereoargenteus ocythous Bangs cinereoargenteus (Schreber) Procyon lotor hirtus Nelson & Goldman lotor Linnaeus Mephitis mephitis hudsonica Richardson avia Bangs Lynx rufus superiorensis Peterson & Downing rufus Schreber

PLAN OF THIS BOOK 19 (1961) for Wisconsin and from Burt (1948) side times two. The dental formula was de- for Upper Michigan. Other symbols are oc- veloped after Baron Georges Cuvier had tab- casionally used for personal observations, ulated comparative formulas for numerous examination of road kills, and other valid mammals to express serial analogy. records. Most “other records” in this study Unless there is solid evidence to identify are designated with open dots. a tooth as one kind or another, and especial- Therefore, for most of the state maps the ly in the problematic insectivores, I prefer to black dots may be compared with the bull’s- avoid use of implied homologies, and prefer eyes or open circles to roughly appraise and to use descriptive symbols such as B for bi- compare the present and earlier distributions. cusp or bicuspid, U for unicusp (above) and Some changes in distribution are delineated unicuspid (below), and so on. In the section with bold dashed lines. (Smaller dashed lines Insectivora further problems with the nomen- already were on the outline maps; they delin- clature of these so-called unicusps are dis- eate drainage divides.) When localities are cussed. I have been advised to at least refer dubious or near one another, I arbitrarily to the first tooth above in shrews as an inci- chose one good locality for mapping and the sor, but am not convinced it is the first inci- others unplotted are italicized in the records sor. An upper fourth premolar is given as P of specimens. The comparison of ranges 4/, and P /4 is the lower analog. past and present show real changes (in which External and cranial measurements and I follow the ornithologists in calling shifts “dis- many other terms used herein are defined in persal”). In some cases, such as many in Wis- the Glossary. Measurements usually are in mil- consin, the change observed is in known range limeters. Dial calipers were used to measure the (where the specimens were recently obtained, skulls. For weights and some linear measure- unseen by Jackson). Usually it is possible to ments either English or metric may be quoted, distinguish one change in geographic range and in instances where the writing is difficult from the other. and jerky from switching back and forth, both The descriptions include cranial and ana- values may be given together. I am persuaded tomical characters, color of pelage, dental for- to provide both values, because of both general mula, size and other known traits. Terms for and scientific readers. Appendix B at the end of color are from common lay use unless quoted this book provides a conversion table for En- verbatim from studies of other workers. Terms glish and Metric Measurements. such as narrow, or narrower zygomata, or elon- Statistics given, unless cited from anoth- gate, and such qualitative generalities are most er worker, will provide individual measure- useful when actual specimens are examined. ments, or if the number (N) is practical the The drawings and photographs of skulls should mean usually will be calculated. The observed be consulted. The finest set of mammal skull ranges and standard deviation are often list- drawings I have seen are in Hall (1981) or Hall ed in the tables. and Kelson (1959). After each account there is Additional Dental formulas used herein are the se- Natural History, including one or more works quence of incisors, canines, premolars, and cited and otherwise deemed appropriate. molars, written for humans (as an example) There is no intent to be judgmental or com- as DF= I 2/2, C 1/1, P 2/2, M 3/3 = 32. prehensive in so doing, some fine works may The premolars 1-2 are not the first and sec- be left out. Finally, geographic variation is ond ones; they are the posterior third and described and discussed. fourth premolars because in evolution the In Specimens Examined, counties are list- anterior premolars are first lost. The total 32 ed alphabetically for easy reference. The coun- is a product of the summed teeth on either ty names and their locations are shown in the

20 THE WILD MAMMALS OF WISCONSIN aforementioned map of counties and type FREE EXPRESSION IN TAXONOMY localities. In addition to the specimens I have examined from the UW-SP collection, I have Some mammalogists may find some of my examined hundreds of University of Wiscon- divergences from established taxonomy dis- sin—Madison museum specimens (UW) doc- concerting, I would like to make clear that umented by Jackson (1961). To preserve the the taxonomy one uses may express a view aforementioned rough comparison between based upon a biological rationale. Darwin the distributional ranges of his day and mine (1859) once said, “Naturalists differ most seldom did I blacken up the circles on the maps widely in determining what characters are of used to represent his old records, even after I generic value.” Every species is distinct, and had seen them, or list those specimens with it often becomes subjective to recognize long my specimens examined. They are listed in phylogenetic separation (i.e., generic diver- his book. With Frank Iwen’s (Univ. Wiscon- gence) from closely resembling (i.e., evolution- sin Madison, Museum of Zoology) help, I tried arily related) species. In Wisconsin six mam- to examine important UW specimens accu- mals present dilemmas: (Microsorex and Sor- mulated since 1961. Some specimens exam- ex, Eutamias and Tamias, Pitymys and Mi- ined by me in the University of Minnesota crotus, Mustela and Neovison, Lontra and Bell Museum are listed as UMinn., in the Illi- Lutra, Lynx and Felis). Some add Puma and nois Museum of Natural History collection as Felis and Bison and Bos. Nomenclature is a UI, and in the U.S. National Museum as system based on rules and regulations com- USNM. Dr. Scott Craven permitted me to piled and published in the International Code examine a useful collection in the Wildlife of Zoological Nomenclature, revised and Ecology Laboratories in Madison (UW-Wild. amended by an International Commission. Ecol.). Although some state accounts today The Code provides for the free expression of omit precise localities taken from specimen diverse classification. Alternative expression labels, there are important reasons to list is welcome. Opinions may be consigned to them. Other taxonomists then learn about the synonymy by other workers hopefully by pre- specimens, which they may borrow for their senting new evidence.* research. If the specimens become lost or destroyed, too often the case in museum collections, the known occurrence is document- PRESERVATION OF SPECIMEN ed. Re-examination by future taxonomists may COLLECTIONS AND HABITATS uncover errors in identification and lead to geographic range corrections. Against these When wild mammals are studied and carcass- positives stand two negatives: the work of list- es obtained or salvaged, voucher specimens ing and the expense of publishing. Here, the should be made and preserved. Students mak- listing was accomplished, but the cost of pub- ing a study should do it themselves as a nec- lication was hard to defend. I followed a middle road; specific localities were uncommon * The interested reader is referred to an excellent and and marginal records. recent synopsis on the subject of mammalian The Literature Cited and Bibliography taxonomy edited by Wilson and Reeder (1993, include some references used by Jackson 2005). For higher categories, Simpson (1945) and (1961), some overlooked therein, many pa- McKenna and Bell (1997) are recommended. Highly pers cited herein, and some worthy but not recommended is the new The Smithsonian Book of Mammals, by the same Wilson with Sue Ruff fully discussed due to the length and expense. (1999) and a host of contributors. A revised Wild To great extent it is bibliographic since Jack- mammals of North America updating Chapman son’s publication. and Feldhamer (1982) has appeared.

PLAN OF THIS BOOK 21 essary part of that study. Although difficult, Table Plan-2Plan-2. The UW-SP Mammal Collection, the time consuming, and requiring skill, invaluable Wisconsin specimens. The Museum contains specimens pre- vouchers should be deposited in a permanent served from nearby states of the several species not in this collection. It contains specimens of Myotis sodalis, Nycti- repository, i.e., a museum collection. Speci- ceius humeralis, Gulo gulo, Spilogale putorius, Cervus ela- mens should be well made, including at the phus, Bison bison, and Rangifer tarandus. least a skin and skull with labels (Appendix C). Today frozen tissues and soft tissues pre- Order Families Genera Species served in alcohol often are preserved. Perma- Marsupialia 1 1 1 nent field notes should be kept with the spec- Insectivora 2 5 8 imens, and all information obtained should be Chiroptera 1 6 7 Lagomorpha 1 2 3 accurate, complete as possible, and meant for Rodentia 6 21 27 permanent record. If there is no way to pre- Carnivora 5 11 17 serve a specimen, even by putting it in a jar of Artiodactyla 1 2 2 alcohol, or saving the skull or teeth of a de- composed carcass, the carcass may be turned over to a mammalogist. That lays a burden on Table Plan-3. Wisconsin specimens at the UW-Madi- someone else to make up the specimen. son Museum. Archaeological bison and the introduced stone Most of the work herein is based on col- marten are included. lections of mammals gathered in the last quar- Order Families Genera Species ter century. Such collections are priceless. Wisconsin specimens are found in several state Marsupialia 1 1 1 Insectivora 2 5 7 collections and in several public museums Chiroptera 1 5 7 outside Wisconsin, but I have not attempted Lagomorpha 1 2 3 to document every specimen. The Mammal Rodentia 6 21 27 Collection at the University of Wisconsin- Carnivora 5 11 19 Stevens Point Museum of Natural History is Artiodactyla 2 3 3 one of several large vertebrate and inverte- brate museum collections. The Museum, a have been made or will be for the following division of the University of Wisconsin— species: Sorex (Microsorex) hoyi, Tamias Stevens Point, is supposed a responsibility of striatus, Sciurus carolinensis, Castor ca- the University of Wisconsin System. nadensis, Microtus (= Pitymys) ochrogaster, The Museum has been the foremost uni- Microtus (= Pitymys) pinetorum), Peromy- versity museum in the state in research col- scus maniculatus, Urocyon cinereoargen- lections, exhibits, and museum activities. The teus, and lesser changes for others. Museum of Zoology at the Madison campus has about as many or more mammal specimens, mostly representing earlier collections. COUNTIES AND TYPE LOCALITIES The two collections taken together provide a respectable repository of Wisconsin mammals The counties with their names are shown on (Tables Plan 2-3). Field notes of many stu- the accompanying map. The records of oc- dents are preserved, and the Department of currence of mammals studied in this book are Natural Resources has provided some vouch- mapped for the counties, which are listed al- er specimens. Some specimens of endangered phabetically. Because a boreal-southern divi- species have accumulated in the collections sion of mammals is important, it is good to throughout the years. know northern counties from southern. The On the basis of the museum specimens, northern ones are often toward the beginning significant taxonomic changes, since l961, of the alphabet (such as Ashland, Burnett,

22 THE WILD MAMMALS OF WISCONSIN Bayfield). Exceptions such as Washburn, Saw- tion where a mammal specimen was collected yer, Iron and Vilas are quickly learned. The and subsequently chosen to represent a sub- Door Peninsula, important for the geography species or species of mammal named formally of Wisconsin mammals, is primarily Door in a scientific published work. Other specimens County. The populous southeastern counties from that same locality are called topotypes, are known by big cities of the same name and they, with or without the type (= holotype), (Racine, Waukesha, and Milwaukee). In cen- are important for taxonomic comparisons of one tral Wisconsin is a square of three counties named kind with another, to associate speci- (Marathon, Wood and Portage) where much mens from other places in the same kinds. of the University of Wisconsin-Stevens Point Most but not all named kinds from Wis- mammal collection was collected. Similarly, consin are valid names in use today. All the the Madison collection is mostly from Dane, names based on types collected at our type Columbia and Sauk counties. localities are available for future taxonomic For convenience, the 17 type localities of opinions. All are cited in the Synonymies that mammals named from Wisconsin are plotted head up the species and subspecies accounts on the county map. A type locality is the loca- beyond. The type localities are as follows:

Map showing counties and type localities.

PLAN OF THIS BOOK 23 Ashland County — Basswood Island, Apos- Jackson (1961) was omitted from his syn- tle Islands: Canis latrans thamnos Jack- onymy, and also by Hall and Kelson son 1949. (1959) and others. The “lead region” of Clark County — Withee: Sorex palustris the Mississippi River is not much like Dane hydrobadistes Jackson 1926. County; in any case, the subfossil elk was Worden Township: Pitymys pinetorum considered an extinct fossil species. Nev- schmidti Jackson 1941. ertheless, it is an available synonym for Columbia County — West Point Township: the American elk. Reithrodontomys megalotis pectoralis Oconto County — Lakewood: Microsorex Hanson 1944. hoyi intervectus Jackson 1925. Door County — Peninsula State Park: Tamias Oneida County — Crescent Lake: Eutamias striatus doorsiensis Long 1971. Jessen’s minimus jacksoni Howell 1925. meadow on Swenson Road, Washington Outagamie County — Near Appleton: Felis Island: Synaptomys cooperi jesseni Long concolor schorgeri Jackson 1955. 1987. [Honoring Tom Jessen, type tak- Racine County — Racine: Sorex hoyi Baird en nearby on his brother’s meadow.] 1858 (= Sorex (Microsorex) hoyi). Prob- Douglas County — Lake Superior, probably ably collected somewhere northward of Valley of the Brule River near Solon Springs, Racine (Long, 1972a). Bruleand Iron rivers. Not Minnesota as re- Racine: Arvicola austerus Le Conte 1853. ported, because Geomys does not occur on Richland County — Lone Rock: Geomys the Minnesota shore or near it. Mus sacca- bursarius wisconsinensis Jackson 1957. tus Mitchill 1821. New York Med. Reposi- Rock County — Milton, 4 mi. E: Taxidea tory 6:249 (=Geomys bursarius). taxus jacksoni Schantz 1946. Grant County — Platteville: Urocyon cinere- No exact locality known except Wisconsin. — oargenteus ocythous Bangs 1899. Cariacus wisconsinensis Belitz 1919 (= Iowa County — Blue Mounds: Cervus whit- Odocoileus virginianus, see Jackson, neyi Allen 1876. Blue Mounds listed by 1961).

Rhyme to Louis Agassiz:

And he wandered away and away, with Nature the dear old Nurse, Who sang to him night and day the songs of the universe, And when his way seemed long and his heart began to fail, She sang a more wonderful song, Or told a more wonderful tale. — Henry Wadsworth Longfellow

24 THE WILD MAMMALS OF WISCONSIN EARLY MAMMALOGY AND NOW to region, and therefore the economics of fur not only affected the history of the American What is mammalogy? It is the study of four- West, but it affected the very early history of limbed furry animals secretive and mostly Canada and the United States. Fur also pro- nocturnal. They climb trees or rocks, dig in foundly affected the destinies of several Indi- the ground a little, and are adapted to our an nations especially along the Mississippi grasslands, forests or forest edge. Some spe- River, and in Wisconsin and Upper Michigan cies run or hop, and others can even fly. They (see Gilman, 1974). Eventually the city of Sault feed on seeds and nuts, sometimes on fruit or Ste.-Marie was settled at St. Mary’s Narrows, grasses, or instead may specialize on insects at the east end of Lake Superior. or flesh. They may breed in the spring or fall, Explorers such as Etienne Brule, Pierre sometimes throughout summer, and a few Radisson, M. Chouart Sieur des Groseilleiers, may breed in winter. Usually the young are Perrot, Jean Nicolet, Father Jaques Marque- born blind, naked and helpless, but some are tte, Louis Joliet, and Daniel G. Dulhut explored more precocial. They nurse, grow, disperse, what is now Wisconsin, establishing schools, and try to carry out their life activities faced churches and fur trading posts throughout the with mortality from predators and diseases. region, from Madeleine Island in the Apostles People observed mammals in what is now to Green Bay eastward and Prairie du Chien Wisconsin over 600 years ago, probably as and La Crosse southwestward. The French early as 9,000-11,000 years ago, the time eventually established settlements along the en- the great Pleistocene mammoths died out. tire length of the Mississippi River. These ac- Fishing and hunting were pragmatic activities tivities of developing a fur trade, and eventual- of native Americans. There were myths and ly of copper mining, fishing and farming all superstitions about mammals. Their lore can- began in the early 1600’s. not be regarded as science, even though our After the French were defeated at Que- science is a changing and imperfect percep- bec in 1759, giving up their American pos- tion of nature (no matter how elegant we be- sessions in the treaty of 1763, the British ruled lieve that it is), we who practice and see our as best they could over the wilderness posts results in retrospect. Retrospection is possi- and the lucrative fur trade, until 1783. Then, ble because of publications, books, journals the Treaty of Paris ceded American lands to and other written records. Publication and the United States. Nevertheless, French, Brit- testable science began in Wisconsin with the ish, and Indians, with the help of some Amer- Caucasians (French, English, Americans). icans, continued shipping furs to European The French established schools, trading markets. The Americans were kept out by the posts, and churches in this region about as British troops until about 1815. Eventually early as the Pilgrims landed at Plymouth Rock. the fur trade and other activities were assumed After the early 1500’s, the active and curious by the Americans, meaning by this term the French explored central Canada and our land people from the United States. By 1836, via the mighty Ottawa River, leading west- Wisconsin was a territory. Its southern lands ward from Quebec. Jean Nicolet arrived at were settled by farmers who early on moved Green Bay in 1643, and he was not the first. along the Lake Michigan shore to French-es- Some coureurs de bois arrived as early as tablished Green Bay and the Door Peninsula 1662 (Gilman, 1974). Most of the fur trap- (named after the strait called Porte des morts- ping and in early years even the transport to or Door of Death). Lumber was harvested in Quebec of the furs were carried out by Native the center of the region, shipped down the Americans. The various tribes were often at Wisconsin River, and eventually lumbering war with one another, displaced from region yielded great quantities of lumber from the

EARLY MAMMALOGY AND NOW 25 tall white pines and spruce of the North ican government-sponsored expeditions to Woods. Lead and copper were mined in the explore the Michigan Territory. His explora- southwest of Wisconsin. Forest fires were tions circled from Sault Ste.-Marie to Grand commonplace. The infertile, sandy savannas Marais, to Pictured Rocks, and westward of the center were eventually settled by farm- along Lake Superior to the Apostle Islands ers. Swamps and marshes were drained, for- (in what is now Wisconsin), down the St. Croix ests cleared, and towns and roads established. and Mississippi rivers as far as Galena, Illi- Thus, there were, in what is now Wis- nois, and in Wisconsin, Prairie du Chien, then consin, several Indian nations, the French, traveling up the Wisconsin River to Portage, British, Canadian, and American govern- northeastward to Green Bay, and back north ments. Even Spanish soldiers visited in the to Sault Ste.-Marie. He made casual observa- southwest parts of this land, after moving up tions (1834) of bears and deer killed, sight- the Mississippi River. For a while the region ings of wolf, porcupine, and in Wisconsin a was known as the Michigan territory. red fox, beaver and “jerboa” (probably a jump- Wisconsin’s borders with Michigan were ing mouse). This mouse was from Madeleine established by a series of political negotiations. Island (Mossman, 1992, 1994). Today there are several Indian nations with On his expeditions Schoolcraft was ac- more or less authority over the northlands. companied by a bona fide scientist named S. The earliest French civilization was more ac- L. Mitchill. Mitchill wrote, “Description of two curately a French-Indian civilization, with mammiferous animals of North America”, mixed culture and free intermarriage. Al- Medical Repository 21: 248-250, 1821, and though these races were amicable, there was “Detection of a mistake into which natural- always trouble between Sioux and Chippewa, ists have been led, in relation to Mus bursar- Chippewa and Huron, etc. Basically the en- ius, or pouched rat of Canada”, Amer. J. tire region had developed in one way or an- Science, 4: 183-185, 1821. other primarily by the economics of fur, until In these early years Wisconsin was visited after statehood, after which time the region by two of the world’s greatest intellectuals. Both was settled by farmers, lumbermen, miners, were greatly interested in nature, and only their businessmen, immigrants, and descendants. brief tenure in Wisconsin keeps them from Although scientists in those early years being recognized as Wisconsin’s greatest nat- faced formidable challenges, and spent much uralists. Both were primarily writers and real- effort in survival, the French scientists and some ist philosophers, but they were amateur natu- others, too, named and described animals from ralists besides, with strong interests in making many of the furs obtained in the fur trade at plant specimens. John Muir (1913) arrived St. Louis and Hudson Bay. None of them is from Scotland in the 1840’s and was raised known to have come from Wisconsin. The with hard work on a farm at Montello. He left badger type may have (see Long, 1972) be- Wisconsin and became an eloquent preserva- cause it was a St. Louis pelt supposedly taken tionist of nature, so elegant in prose to be al- in “les pays des Esquimaux” which generally most biblical, so outraged and stately he re- was French-Canadian fur territory. French ex- sembled a biblical prophet. Never has there plorers such as Joliet and Nicolet made gener- been another like him, and nobody from Wis- al observations on wild mammals in their ac- consin is really educated if he knows nothing counts of the regions explored. Joliet [“Louis of the lyrical and sacred environmental scrip- Jolliet”] described a mountain lion in what is tures that he wrote. Muir described the nest now Grant County in 1673 (Jackson, 1961). and a hibernating specimen of the thirteen- With the fur trade flourishing in the early lined ground squirrel Spermophilus tridecem- 1800’s, H. R. Schoolcraft made several Amer- lineatus. Quite interesting was the observed

26 THE WILD MAMMALS OF WISCONSIN attack (about 1846) of a shrike, even following a ground squirrel into its burrow. The other intellectual giant was Henry David Thoreau, who briefly visited Wisconsin, and paid more attention to mammals than did Muir. Accompanied by Horace Mann, Jr., the transcendental realist Thoreau came by steamboat up the Mississippi River hoping eventually to write up an account of the Great River and the Volga-like towns here and there along the river’s steamboat route. These included Prairie du Chien, La Crosse, Red Wing, Min- nesota, and others. Some little towns are still isolated along that river today, although a highway and railroad now extend along the river shore. Thoreau unfortunately died before he wrote the book, although publishing a few articles. His notes are preserved at Harvard Uni- versity. His favorite mammals were the thirteen-lined ground squirrel Spermophilus tridecemlineatus and the pocket gopher Geomys bursarius (Sweetland, 1962), which is indeed John Muir. Great naturalist-philosopher. Raised near a more interesting mammal than most people Montello. Photo by W. Dasselville, about 1909. Courtesy of realize. The woodcut of Thoreau was made the Sierra Club. after he returned to New England from Wis- consin, shortly before he died. Several works on the Wisconsin mammal fauna appeared in the late 1800’s. I. A. Lapham (1852-1853) and Moses Strong (1883) wrote lists of the mammals. As early as 1853, Dr. P. R. Hoy wrote a paper on the thirteen-lined ground squirrel. Hoy sent numerous specimens to Spencer Fullerton Baird at the Smithsonian Institution. He actually named with Kennicott a kind of deer mouse found in Wisconsin, named bairdii, and had a shrew Microsorex hoyi (=Sorex (Microsorex) hoyi) named after him by Baird. In 1882, Hoy wrote a brief account of the mammalia of Wiscon- sin. H. Gillman (1873) wrote an early work on the caribou near Lake Superior. Other naturalists at this time were Charles F. Carr, John Clark, Henry Nehrling, and the two Kumleins (Thure, died 1888; Ludwig, 1880’s). One of the earliest mammalogists living in Henry David Thoreau. Naturalist and transcendental Wisconsin, and, indeed, the first woman mam- philosopher. after returning home from Wisconsin Mississippi malogist in America, was Martha Maxwell. Fa- River trip. Woodcut.

EARLY MAMMALOGY AND NOW 27 mous for her Colorado exhibition at the 1876 Illinois to boreal Wisconsin. As comprehensive Centennial celebration in Philadelphia, Maxwell as Jackson’s work on Wisconsin was in 1961, was mainly a self-taught collector and taxider- no one should overlook Cory’s practical dis- mist of birds and mammals. When living in Bara- cussion on any Wisconsin mammal. boo she married, had a daughter, and got work Dr. Hartley H. T. Jackson (1881-1976) at the Baraboo Collegiate Institute. Her work was born in Milton and studied at Milton Col- was to assist in the preparation of mounted birds lege. He worked for his masters degree at and mammals for the Zoology Department, a U.W.-Madison and obtained a doctorate from practice prevalent at state schools at least until George Washington University in 1914. His 1906. When she left Wisconsin for Colorado undergraduate teacher was Ludwig Kumlein she had learned to observe, study, collect, and (see Robbins, 1991). His first paper on Wis- prepare specimens mounted in natural positions, consin mammals was published in 1903. In including large mammals. She corresponded 1910, he joined the U.S. Biological Survey, with Joseph Henry, Secretary at the Smithoso- and although he spent most of his life outside nian, and also with Robert Ridgway and Elliot Wisconsin he may be the greatest naturalist Coues. A screech owl that she collected was the state has produced. He was instrumental named after her by Ridgway. She also confirmed in founding the American Society of Mam- the existence of th black-footed ferret described malogists, and was a longtime editor for the earlier by Audubon. She attended college in Wis- Journal of Mammalogy. He was a meticulous consin (Lawrence University) at a time when scholar, and wrote a classic taxonomic mono- women were generally not welcome in college. graph on the shrews of North America. He Like Muir she subsequently made her fame in a also collaborated with Stanley Young on western state. Her collections of specimens books such as The Clever Coyote and finally apparently are all lost (Stein, 1996), unless a wrote the Mammals of Wisconsin (1961). few are in the United States National Museum Aldo Leopold (1899-1948), born a cen- (Natural History). tury ago, was a forester acknowledged as a About 1900, workers from the United great naturalist-philosopher. His books (e.g., States Biological Survey, under the direction Sand County Almanac, Round River) ex- of Dr. C. Hart Merriam in the Smithsonian pound a “land ethic” philosophy. He provid- Institution, began to collect specimens in Wis- ed classical foundations for game manage- consin. Two young Wisconsin naturalists, Ned ment, recognition of the ecosystem, and an Hollister and Hartley H. T. Jackson, learned appreciation of the role of predation in regu- about the Smithsonian and actually got jobs lating prey populations. For Wisconsin he with the Biological Survey. Classmates at Mil- published on wolves, cottontails, and the “kill- ton College, and fellow workers in the Sur- ing technique” of the [long-tailed] weasel. vey, it is not surprising that early works on In the 1940’s, bat banding was popular Wisconsin were produced by both of them. in the caves of Wisconsin. Pioneer bat band- Hollister (1910) wrote an early check-list, and ing carried out by professors and students at in his youth Jackson (1908) wrote a prelimi- Madison was done at Eagle, John Gray, and nary listing of Wisconsin mammals. nearby caves by John Emlen and William El- Two scholars who wrote several early pa- der. James Beer, Frederick Greeley, and Ar- pers on Wisconsin mammals at the turn of the nold Jackson banded bats in southwest Wis- 20th century were W. E. Snyder (1902) and consin, at Lac du Flambeau and sites near H. L. Ward (1907, 1908, 1911). In 1912, C. Hurley. Wayne Davis, who reviewed the bat B. Cory wrote the “Mammals of Wisconsin accounts herein, and W. Z. Lidicker, gradu- and Illinois”, which is a comprehensive and ate students at the University of Illinois, stud- well-written book on mammals from southern ied bats from southwest Wisconsin.

28 THE WILD MAMMALS OF WISCONSIN L. B. Keith, Department of Wildlife Ecolo- counties have set aside county parks not en- gy at Madison, and his associates made nu- tirely developed for recreation. Since 1960, merous contributions on the cycles of hares The Nature Conservancy has preserved or and lynx. A. W. Schorger, Wildlife Ecology, encouraged the preservation of about 40,000 studied the newspapers and county histories acres of Wisconsin wilderness. Robbins to compile histories of several mammals and (1991:32) has mapped the national wildlife birds of Wisconsin, and he made a list of the refuges, state wildlife areas, and major re- mammals of Dane County. Since then we have search stations of the state. There are about seen quite a lot of ecological work by the De- 194 scientific areas preserved by legislative partment of Natural Resources (formerly the action by 1951. Much of this conservation Wisconsin Department of Conservation) includ- stems from the teaching, eloquent philoso- ing that of Jim Hale, W. E. Scott, G. J. Knud- phy and sincere efforts of Aldo Leopold and sen, N. R. Barger, and others. Today there his students. Other important naturalists of are at least three departments of the DNR di- the past included John Muir, William T. Hor- rectly related to mammalogy, the Bureau of naday, Ernest Thompson Seton, and even Research, the Bureau of Wildlife Management, President Theodore Roosevelt, who instilled and in recent years an important new Bureau by eloquent writings in those days a wide- of Endangered Resources. One person who spread caring for nature. They sparked nec- may never have visited Wisconsin at all, but essary action to preserve our natural heritage. who mapped and classified its mammals of- Some Basic Procedures. In the earliest ten, with authority and in a broad scheme of years American mammals were examined in classification was E. Raymond Hall, University hand after collecting them with gun or steel of Kansas, my former teacher. In his The Mam- trap, or when carried in by native Americans. mals of North America (1959, with K. R. They were mounted usually in a fierce demean- Kelson, and 1981) he summarized the litera- or, in some sort of natural position, or pre- ture including some of his own papers treating pared as a pelt. The skull was often discarded, Wisconsin specimens, e.g., microtines, wea- or the skull might have been left in the stuffed sels, lagomorphs and others, and mapped ev- head. Invention of the snap trap, effective for ery species in the state not only for Wisconsin catching small mammals and cheap to manu- but providing maps of their geographic ranges facture in quantity, led to huge collections of throughout North America. [I relied on them the mice and rats in our museums, with an heavily for the North American maps herein.] occasional shrew or ground squirrel. The spe- Today, in spite of great population growth cially designed museum special trap, which is and intensive land use, Wisconsin still main- an expensive trap, strikes the small mammal tains a rich diversity of natural communities on the neck or back and seldom crushes the under federal and state protection. Since skull. This development, following the tech- 1971, the state DNR has provided increasing nique discovered of using dermestid beetle lar- attention to endangered species. Many stud- vae to thoroughly clean small mammal skulls, ies on wild mammals are done cooperatively led to better skull collections—useful in taxo- with state agencies in nearby Upper Michi- nomic work. “Skin and skull” became the spec- gan, Minnesota, and with the Federal Gov- imen of modern mammalogy, with practices ernment. Two national forests (Chequamegon instilled in scientifically trained professionals by and Nicolet) were established between 1928 the likes of Joseph Grinnell and Spencer Ful- and 1933. The DNR established great re- lerton Baird, and eventually E. Raymond Hall. serves for wildlife at Horicon, Crex Meadows They all stressed keeping permanent well-writ- in Burnett County, the Mead Wildlife Area in ten field notes on the ecology of the mam- central Wisconsin, and in other places. Some mals. Mole traps, pocket gopher traps, and

EARLY MAMMALOGY AND NOW 29 the Japanese mist net made of fine threads cially of rare carcasses obtained for deposition (invisible to flying bats) led to better collections in a museum. Unfortunately, too often the sal- of some heretofore obscure mammals. The use vaging of specimens does not happen in Wis- of empty cans sunk into the ground flush with consin. Common sense and tact are impor- the surface of it, often called a pit trap or can tant for mammalogists, who should not jeop- trap, has led to easy collection of shrews (Han- ardize their opportunities to study animals by dley and Kalko, 1993). Drift fences designed alienating people who object to cruelty and to funnel mammals into the traps increase the killing of animals. After all, it is in the best in- catch. Once dug in, the cans are easy to check terests of the wild animal for workers to know over a period of days or weeks. Today mam- its biology for proper management and its pres- mal characters also are obtained from blood ervation. A policy statement (Acceptable Field sera, DNA, isozymes, the post-cranial skele- Methods in Mammalogy. . . . 1987) of the ton and other body parts. Live traps permit American Society of Mammalogists has been marked mammals to be released and re-cap- issued on this subject to all members. tured. Small radio transmitters allow some The American Society of Mammalogists’ mammals to be followed wherever they wan- publications Journal of Mammalogy, Mam- der. Keen observation of animal and habitat malian Species, and Special Publications, always was important, and a fine camera and is an important resource. The annual meet- pair of binoculars are often useful in research. ings and constant appraisal of the role in so- Field collecting has its difficulties because ciety by mammalogists lead to a respectable for nocturnal mammals the traps must be set and scientific enterprise. The Journal of Wild- on one day and checked on the morning of life Management, published since 1937, also the next. Preparation of the specimens, and contains many articles on mammals. securing both skins and skulls from loss or damage, take much time and effort. Traps must be set so that mammals will find them, which requires some knowledge (“lore”) about the secretive behavior of the mammal, and of the mammal’s “sign” (e.g., tracks, hairs, runways, burrows and scats). Hall, in Hall and Kelson (1959), and other references describe the steps for making study skins. This procedure is given in Appendix C. Proper state permits must be obtained from state natural resources agencies to collect mammals. These rarely are available for amateurs, and are reserved for professional mammalogists to further scientific investigations. Skins and skulls in personal collections are often illegal. Rare and endangered species are seldom permitted to be collected, and seldom are game animals allowed to be killed out of season unless a rationale is approved. Some natural history museums should be selected for final deposition of specimens, even of carcasses found. Hartley H. T. Jackson. U. S. Biological Survey, Most mammalogical studies would benefit by Washington, D. C. Born in Wisconsin, wrote on mammals the preparation of voucher specimens espe- more than 60 years. 1938.

30 THE WILD MAMMALS OF WISCONSIN ENVIRONMENTS AND These wetlands provide homes, feeding and ECOLOGY OF MAMMALS breeding areas to many kinds, such as jumping mice, shrews, bog lemmings, beavers, GENERAL ECOLOGY OF muskrats, mink, and otters. Great rivers (see WISCONSIN MAMMALS section on zoogeography) restrict some mammals mostly from dispersing eastward, but the Prairie, Forest Edge, and Forest sandy floodplains were routes of dispersal for From The prairies — ”... My heart swells while the the pocket gopher. Rocky bluffs with caves dilated sight takes in the encircling vastness... The and mines provide shelter for millions of bats. clouds sweep over with their shadows and beneath, Basically, the chief contingents of mam- the surface rolls and fluctuates to the eye... Breezes mals are boreal, i.e., northern (e.g., north- from the South who toss the golden and the flame- ern flying squirrel, Canada lynx, moose) and like flowers and pass the prairie hawk that, poised southern (e.g., prairie (= eastern) mole, least on high, flaps his wings, yet moves not. From shrew, opossum). Boreal species have occu- Entrance to a Wood — “Enter this wild wood and pied the North Woods for centuries, and have view the haunts of nature. The calm shade shall dispersed southward even somewhat beyond bring a kindred calm, and the sweet breeze that the limits of present day boreal forests. South- makes the green leaves dance... of green and ern invaders from the deciduous woodlands, stirring branches alive and musical with birds... The forest edge, and prairies in the south have squirrel, with raised paws and form erect, chirps moved northward into the northern forests. merrily. Throngs of insects in the shade dance in By way of sand dunes and beaches, wetlands, the warm beam that waked them into life... Even wet prairies, and savannas the dense, shady the green trees partake as they bend to the soft forests were penetrated by many sun-loving winds, the sun from the blue sky... From Forest prairie and forest-edge mammals. This inva- Hymn — “All these fair ranks of trees. They, in thy sion occurred even before Europeans and sun, budded and shook their green leaves in thy Americans arrived. After human settlement, breeze, and shot toward heaven... When I think of the clearing of forests by farmers and lum- the great miracle that still goes on, in silence around bermen and tree removal for road rights-of- me — the perpetual work of thy creation, finished, way encouraged the northward dispersion of yet renewed forever.” — William Cullen Bryant. southern species. But at the same time numerous boreal species, and some grassland The southern prairie soils, rich in organic kinds such as the least shrew (Cryptotis par- matter, support dense grass and forbs; the va), were disappearing probably because of friable and well-drained soils shelter numer- agricultural practices. Chemicals used in the ous burrowing mammals, especially seed, environment may have decimated and eradi- grass and insect eaters. The extensive and cated local populations of some mammals. often continuous northern and eastern for- Until there was sensible management of game ests of Wisconsin provide mast crops, tree animals and furbearers, some mammals were canopies useful to arboreal mammals, nest eradicated by unregulated hunting and trap- cavities and loose bark for their homes, and ping. A few such species have been reintro- correlated ecological factors such as less snow duced. Habitat destruction and agricultural cover, more leaf litter and less wind chill. The practices in Wisconsin are the greatest threats southern hardwoods provide mast and cover, to Wisconsin mammals today, although the and in the clearings the forest edge mammals latter is often beneficial to animals that need thrive. Rivers and lakes are homes for many openings and forest edge. semi-aquatic mammals, as are the vast There are many positive things that should swamps and countless marshes of Wisconsin. be emphasized regarding wild mammal con-

ENVIRONMENTS AND ECOLOGY OF MAMMALS 31 servation in Wisconsin, and the preservation much to temperature in central Wisconsin. I of mammal environments (many by hunters suppose Rib Mountain may be even warmer and for hunting). Reintroductions of some lost in summer on its high crest than downhill by species, protection of wilderness areas, ac- the Rib River. Central Wisconsin shows more quisition of new refuges, scientific manage- effects of wind chill and deep frost than do ment of harvested mammals, and protection northern habitats near Lake Superior. Much by the rare and endangered species programs greater ecological importance than tempera- have been remarkably successful, to name ture effects should be attributed to the geo- only a few programs. These are primarily the morphology of this state, the major plant com- responsibilities of the Wisconsin Department munities that developed after the recession of of Natural Resources. But recent losses from glaciation, the effects of ecological succession our fauna of some species without notice, the and land use, and the zoogeographical inter- dramatic decline of others, and the relentless actions of invading and retreating mammals. assault on natural habitats by urban sprawl, highway development, and many farmers and builders (in both city and country) should not FORMER GLACIATIONS be allowed to cause further losses of any mammals from our fauna. Even counties and indi- The Pleistocene Epoch, popularly called the viduals must get into the conservation effort. Ice Ages, consisted of four continental glaci- No resting on the laurels of past successes is ations extending as far southward on the sensible. Much proposed land use today is North American continent as central Iowa and detrimental to nature, and many political northeastern Kansas. The epoch lasted about maneuvers nationally and at local levels seem 2 million years, and the climate created most to be intensifying environmental change as of our landforms and soils over vast regions so-called “progress”. In the popular rationale (Dorr and Eschman 1971). of profit and jobs, even a protected nature The glacial stages were not simple advanc- preserve seems for some people merely a es followed by melting and recession, but were reserve. I am advised that at our state level, often complex, consisting of two or more large area land use planning, comprehensive advances with various side lobes from the gla- land management, and ecosystem manage- ciers. Low hills comprised of glacial sediments ment present a sound approach and method (called drift) were deposited by the ice fronts for accomplishing the goals of genuine con- and are identified as lateral or terminal mo- servation. They are all based upon the sci- raines. They mark the extent of the advances ence of ecology. of ice. The moraines can be dated (i.e., aged). Updating ecological information from Interspersed with the four cold glacial peri- Jackson’s treatise (1961) on mammals is nec- ods were three interglacial periods compara- essary particularly in comparing the concepts ble to our present day climate. Although the of his day to those of modern ecology. The southwestern hills of Wisconsin were bypassed fundamental concept in Jackson’s work is the by glaciers repeatedly (giving basis for the singular effect of temperature roughly dividing name “Driftless Area”), the climate was doubt- Wisconsin into three faunal subdivisions, name- less frigid during the glaciations. Where for- ly three of C. Hart Merriam’s life-zones. Jack- ests subsisted at all, they were mostly spruce, son’s perception of temperature was inade- and the mammals in them were boreal. On quate and in some places inaccurate. Further- the snowfields and ice, the mammals were more, the life-zone concept does not work well like those in the Arctic. except in mountainous regions of western Glaciation scoured much of the Wiscon- America. Neither latitude nor elevation relates sin terrain during the Pleistocene Epoch end-

32 THE WILD MAMMALS OF WISCONSIN ing about l0,000 years ago. Just preceding CLIMATE the end of Pleistocene glaciation, late in the Wisconsin glaciation, a huge glacier sprawled The climate is relatively uniform today in Wis- across the northland, lay in the Lake Michi- consin, in average temperature, winter and gan basin, and extended westward across the summer temperature, and precipitation (Table valley of the present day Fox River (Long Env-1). The warm air of summer pushes north- 1974). Earlier glaciations had already planed ward from the south, and it is warmer in the much of what is now southern Wisconsin. southern and central counties with more hot Except for the non-glaciated Driftless Area in days. Winter or summer, the cold air masses southwest Wisconsin and scattered exposed originate in the Canadian Arctic regions and outcrops of rock, the terrain today features usually swing eastward from Minnesota or flow post-glacial phenomena of either the Wiscon- across Upper Michigan and southward along sinan or earlier glaciations. Glacial processes Lake Michigan. Prevailing westerlies bring affected geomorphology almost everywhere. moisture eastward, usually from Minnesota. Two subsequent processes, of course, have There is a mingling of cool Arctic air and south- been at work: (1) stream erosion, and (2) soil ern warm air. As a consequence from temper- formation. The vast glacial outwash sands ature and atmospheric pressure changes, pre- and sands eroded from a sandstone bedrock cipitation increases eastward, and the eastern in central Wisconsin may lead to deep frosts forests have higher humidity. The temperature there in winter and relate to extreme sum- gradients change regularly and gradually north mer heat and drought. Large elongate and to south, as does photoperiod (= day length) interconnected glacial moraines extend and snow cover (with exceptions). The great across the state. These hills are usually cov- freshwater lakes Superior and Michigan mod- ered with woods. Sands, boulders and clays, erate climates near the shores causing the sea- with acid soils, are prevalent. Lying south- sons there to lag. ward of the Wisconsin glaciers, in what is The growing season (100 days in the now Illinois and southern Wisconsin, the flat north, 170 days in the south), based on frost- country scoured by earlier glaciation was left free days, is variable in Wisconsin. Though to nature. The prairies that developed there roughly related to latitude, the season varies had deep, black topsoil, rich in organic ma- from one city to another (Moran and Mor- terial. This has been largely put to the plow gan, 1976), and the extreme numbers are in historic times, but here and there prairie variable and not always as expected. Frost remnants and old field habitats have allowed most of the prairie species of our Wisconsin Table Env-1. Climatic Factors of Southern and North- mammals to survive in spite of farming. A ern Wisconsin (After Curtis, 1959), generally prairie and huge lake called Glacial Lake Wisconsin forest. drained from the terminal moraine near Stevens Point southwestward. It exists even Prairie // Deciduous Forest today as marshes, flowages and drained January mean temp* 15.1 degrees F // 11.40 F marshes from the Buena Vista in Portage July mean temp 71.1 // 68.0 County to the enormous Pettenwell Flowage, Annual mean 44.6 // 41.5 extending southwest toward the Driftless Growing season* 148 days // 126 days Annual precipitation 31.5 // 30.1 in. Area of the hilly and rugged Coulee Country Annual total snowfall* 41 in. // 5l in. in southwestern Wisconsin. Over most of the Number days 0 precip* 90 days // .05 days state is a thin wind blown glacial loess inter- Ave. evaporation (July)* 5-6 in //3-5 in mixed with forest humus and soils derived Warm, days above 68 0F* 60-90 days // 0-60 days from the underlying bedrock. *Apparently significant to mammalian distribution.

ENVIRONMENTS AND ECOLOGY OF MAMMALS 33 penetration of the ground is a variable phe- At dawn of a new millennium there is great nomenon from place to place in Wisconsin, concern that the climate is steadily warming, as is the effect in some places of summer due to so-called “greenhouse effects” of atmo- drought. Both of these seem most severe to spheric pollution, El Nino ocean effects, and the small mammals on the sandy soils, espe- other causes. So far it seems difficult to con- cially of central Wisconsin. Greater snowfall firm such a trend, but one notes the past eight in Upper Michigan and northern Wisconsin or ten winters have been mild, especially 1999- limits frost penetration there, thereby dimin- 2006 (but 2000-2001 was colder). Record high ishing frost-caused mortality of small mam- winter temperatures for three years relate to mals. Snow cover affects distribution, favor- the waters of Lake Michigan that receded sig- ing mammals such as deer that move into the nificantly (although U.S. Army engineers deep coniferous woods, and hares and lynx, dredged out its outlet eastward). Nationally which run easily over the snow. Some tem- there are severe droughts and shortages of perature data are mapped herein (see accom- water; and there seem probable effects on wild- panying Fig.). life (e.g., increasing deer densities). Slight differences in mean temperature throughout Wisconsin do not sharply influence mammal distributions. Rocky outcrops, humus- PHYSIOGRAPHY covered forest soils, sun dappled prairie sod and bare sands, shaded barren woodland soils, To better understand the ecological role of riparian vegetation and wet soils, and other physiography, often called geomorphology, will innumerable local factors provide a diversity reveal interesting and paradoxical effects in of mean temperatures. These allow an inter- Wisconsin. In this author’s experience, in the mingling of boreal and southern species across rugged mountain-basin topography of the the middle of Wisconsin, a grouping that helps Rockies, and even in the flat cuestas of Kan- define the “Tension-Zone.” sas, there is a correlation of natural effects of temperature, climate, vegetation, and animal distribution, all relating to the basic land forms. In Wisconsin, the provinces mapped by Mar- tin (1932) in his classic works derive almost entirely from the single source of underlying bedrock. Indeed, both non-glaciation (in the rugged Driftless area of southwest Wisconsin and as well in the vast swampy marshlands southwest of Wisconsin Rapids) and Pleistocene glaciation have subtly affected the Wisconsin landscape. They have drastically modified the land of granitic rocks of northern Wisconsin showing an almost recent effect (youthful stream development, marshes, lakes). On the Central Plain vast deposits of sand and subsequent sandy soils (drained out of glaciers) are found. The igneous bedrock in the north and the underlying sandstone in the center prima- Winter Severity (includes temperature). Mild to 50, rily create the different physiography. Perhaps Moderate, over 50 is Severe. 1961-90. Bartelt et al. Compare the mammals that range through several prov- Vernon and Marinette. inces only follow certain plant communities,

34 THE WILD MAMMALS OF WISCONSIN and since the land is not tremendously varied 5. Eastern Ridges and Lowlands: This in elevation the pervasisive factor of physiog- region is associated with good soils, flat ter- raphy seems less important than in other states. rain and cuestas with limestone and shale In 1932, Martin described the physiogra- bedrock including a higher, more rugged out- phy, or geomorphology, of Wisconsin as five cropping of the Niagara Formation lying east- provinces. ward along Lake Michigan. 1. Lake Superior Lowland: The good soils All of these physiographical provinces re- there are underlain with ancient pre-Cambrian late somewhat to the distribution of mammals, rocks of igneous and metamorphic structure, except that the mammals range freely through and lie approximately in the basin of Lake one to another depending on the prevailing Superior. vegetation and their mobility. For example, 2. Northern Highland: This province con- moose and lynx enter Wisconsin in the Lake sists of most of boreal Wisconsin, and con- Superior Lowland, but both species range tains swamps, lakes, marshes, and shallow southward and eastward in the boreal forests streams of youthful drainages since the re- of the Northern Highland, and some boreal treat of the Wisconsinan glaciation. Here are mammals range even into the swamps near found boreal forests, probably held over from Necedah, the hills of the Baraboo Range, or the climate of the late Wisconsin. This land the forests along Lake Michigan. There are was not planed off by glaciers as is often Western Upland prairies more or less connect- thought, but is an old peneplain of a great, ed ranging from the low Mississippi Valley and worn-down mountain range, with its hard grassy western “goat prairies” on the unfor- bedrock foundation submerged in seas, but ested ridges, through the openings in the sa- re-emerged from them about 200,000,000 vannas of the Central Plain. These rolling grass- years ago. Nevertheless, the peneplane was lands become intermixed with hardwood for- often scoured and planed by Pleistocene gla- est, creating much forest edge and savanna, ciers from several sources. extending along the Wisconsin River’s south- 3. Central Plain: This consists of sands ern boundary. There the proximity to the and savannas with extremes of heat, drought Grand Prairies Province of Illinois and rich soils and frost, but also includes the unglaciated dating from the early glaciations allow the free swampy, marshy areas southwest of the edge dispersion and deep penetration of prairie flo- of the Wisconsin ice sheet. This entire region ra and fauna into Wisconsin. To the five phys- of some 13,000 mi2 of generally infertile soils iographic provinces I add two important geo- (but blessed by irrigation and now becoming logical features: Wetlands and Islands. useful for peat and cranberries) is entirely Islands. There are numerous islands to- underlain by Cambrian sandstones that bor- day clustered in Lake Michigan (e.g., Big Sum- der the northern igneous bedrock northward mer, St. Martin, Rock and Washington) and and several sandstones and limestone bed- Lake Superior (e.g., the Apostle Islands: rocks eastward, and southward. Madeleine, Outer, and others). Both areas 4. Western Uplands: These are largely were glaciated perhaps even 10,000 years prairies or savanna regions, including impor- ago, so that only in protected places could tant river terraces and lakes of the Mississippi mammals have survived there. Probably these River, and the presence of other deep rivers mammals were boreal (see Pre-historic Mam- (Black, Chippewa, St. Croix, and the lower mals). Today the waters are cold and rough, reaches of the Wisconsin). Rugged bluffs of so that rafting on floating logs and branches sandstone and limestone are found in this is uncommon, and hibernators, inactive in region important to cave-dwelling bats, but winter, cannot cross the winter ice. There- the elevation of these hills is not high. fore, many mammals cannot invade these

ENVIRONMENTS AND ECOLOGY OF MAMMALS 35 isles, but if somehow established there those shrews, snowshoe hares, red-backed voles, species were isolated on the isles. This has meadow voles, bog lemmings, weasels, white- led to some interesting evolution, even in a tailed deer, and even an occasional bobcat, brief 10,000 years (geologically speaking). Canada lynx, or moose. Open marshland Two races have been named from the Door with wet soils near standing water containing Peninsula, a chipmunk Tamias striatus plants such as swamp loosestrife, royal fern, doorsiensis and a bog lemming Synaptomys blue vervain, sedges, rushes, grasses, and such cooperi jesseni. No significant speciation has shrubs as red-osier, dogwoods, willows, Spi- been observed on the more recently uncov- raea, and aspens have mammals including red ered Apostle Islands. Deer mice show micro- foxes, coyotes, weasels, meadow voles, jump- geographic variation on both island groups. ing mice, occasional bog lemmings, Frank- Bats and aquatic mammals easily traverse the lin’s ground squirrels, mink, and otters. Cat- water barriers, and carnivores readily cross tail marsh is an important habitat for masked either water or ice. Humans have introduced shrews (Sorex cinereus), muskrats, jumping some species to some isles (Long 1978c; Long mice (Zapus hudsonius), Franklin’s ground and Long, 1993). Islands are discussed more squirrels, mink, and otters. Southern fens, as fully in the section on Zoogeography. described for Wisconsin by John Curtis, with Wetlands. Neither forest nor prairie can alkaline water, generally contain a few mam- establish itself well when conditions are too mals, such as the Southern bog lemming, and, wet. The “wetlands ecosystem”, if extensive nearby on the brushy shores, white-footed enough, consists of a mosaic of variable mi- mice. Red squirrels (Tamiasciurus hudsoni- crohabitats in association with semi-aquatic cus), gray squirrels (Sciurus carolinensis) or and somewhat aquatic mammals. Recently flying squirrels (Glaucomys volans) may in- and suddenly, Wisconsin wetlands have come habit the nearby trees. Riparian streamsides to the fore, and have been popularly protect- provide important habitat and streamside cor- ed. Mammalogists must study them. Bats, ridors for many kinds of mammals. Wet mead- raccoons, voles, and the semi-aquatic musk- ows, with sweet flag, milkweed, golden rod, rat are found in most of them, and Blarina, sedges, bulrushes, marsh marigold, and tran- deer mice, jumping mice, red-backed voles, sitional plants stand between forest and aquat- tree squirrels and chipmunks inhabit copses ic habitats, so that the mammal fauna is rich, of trees and shrubby areas at many wet local- varied, and usually seral. ities. The wetlands differ in their vegetation, water quality, and wildlife. Generally, the greater the amount of vegetation, the more SOILS mammals are found. Some wetlands, such as Shorelines and Sand beaches rarely provide The topography, about 1,000 feet (= 305 m) habitat for mammals, but even sand dunes, elevation overall, may lack dramatic relief, but with scattered vegetation of peculiar and reg- there are several outstanding features of the ular beach-dune flora, shelter an occasional terrain apart from the vegetation on it. The white-footed mouse and local populations of non-glaciated hilly southwestern part (the cou- the prairie deer mouse Peromyscus manicu- lee country, meaning valley country) is a rug- latus bairdii. Beaches of lake-pebbles and ged region with high wooded slopes and nu- gravel are visited by mammals from shrews merous crevices, ravines and caves. The sand to bears, but have no known permanent res- plain in the center of Wisconsin, although idents, occasionally housing a lodge of bea- heavily wooded and penetrated by streams vers. Tamarack, cedar, spruce, and alder and rivers, is an important geological feature swamps and bogs contain Arctic and masked due to such vast deposits of sand. The lakes

36 THE WILD MAMMALS OF WISCONSIN and marshes that dot the North Woods con- Lake Michigan). Most of northern Wisconsin stitute a vast wetland-boreal wilderness still is overlain with poor acid soils on the uplands not much cleared by human hands. The pale (Iron River, Kennan soils) and reddish brown soils and outcrops of the Niagara Rock For- sandy soils on flat and rolling terrain (Omega mation on the Door Peninsula and isles off- and Vilas soils). These may cover the granitic shore, as well as the isolation and semi-isola- and basaltic Cambrian rocks. In the south are tion of mammal populations there, comprise rich sandy loams (Fayette, Dubuque, Wea, a highly interesting laboratory of recent evo- Warsaw, Dodge, McHenry, and Casco soils). lutionary processes. These prairie soils merge northward into poor- Outcrops of bedrock, such as granite or er sandy soils based on sandstones and silt- limestone, the post-glacial soils and till, sandy stones in the west (Hixton, Norden, Sparta outwash deposits, river bottoms, peatlands soils) and on glacial outwash sands in the cen- and sandy loam prairies are diverse habitats ter (Plainfield, Oshtemo soils). The Fox and necessary for particular mammals in Wiscon- Wolf river valleys are limey and sandy soils sin. Most of the soils are acidic, often infer- overlying drift and limestone strata, which tile, and range from very dry to very wet. They often are conspicuous outcrops and bluffs in are classified as follows: Door County. A surprisingly large total area 1. Coarse, bouldery acid soils derived of in Wisconsin consists of low, flooded soils igneous and metamorphic rocks of the North- (Elba, Poygan, Newton, and Arenzille) com- ern Highland (see above). prised of peat and muck (e.g., Houghton soil). 2. Acidic and infertile sands from sand- The prairie mammals, such as Spermo- stones and glacial outwash. philus tridecemlineatus and Peromyscus m. 3. Fine-grained alkaline soils derived most- bairdii, both of grassland and scattered brushy ly from dolomite (limestone). copses, inhabit the southern loams and range The former glaciers produced clay, sand, onto many accessible open sandy soils north- gravels, silt and indirectly (with accumulated ward. The boreal mammals, such as Clethri- mossy vegetation) deposits of peat. Reddish onomys gapperi, Glaucomys sabrinus, and lake clays and sandy loams were produced, Myotis lucifugus occupy mainly the forested mostly in eastern Wisconsin along Lake Mich- gravels and coarse, sandy acidic soils derived igan and Lake Winnebago, but also along the of igneous and metamorphic rocks. They Lake Superior shore. Sands were deposited make up much of the terrain of the northern up to 60 feet (18.3 m) depth on the Central counties. Aquatic and marsh-dwelling mam- Plain, and are widespread across six or seven mals thrive on the low, flooded soils in peat counties (a broad band often exceeding 100 bogs and other wetland communities. Out- miles (= 161 km) width). Another band of sand crops of limestone (dolomite), granite, and extends along the St. Croix River northeast rough country generally, are habitats of mam- to nearly as far north as Bayfield. A deep (up mals such as chipmunks and long-tailed deer to 16 feet), so-called silty loess lies along the mice. Both large and small caves in the sand- Mississippi River in southwest Wisconsin. stone or limestone outcrops and river bluffs Loess is also found in Dane and several ad- in southwestern and western Wisconsin are joining counties (Zimmerman, 1991). homes to thousands of bats. A quick overview (Hole et al., 1966) of There may be remarkable differences be- Wisconsin soils, named generally from geo- tween the humus and topsoil of deciduous for- graphic names, mentions some clayey soils est and coniferous forest stands. Pine needles concentrated along the shores of the Great cause soils to be remarkably acidic. Deciduous Lakes (Hibbing, Ontonagan soils near Lake leaves produce a deep leaf litter with less acid- Superior; Manawa and Kewaunee soils near ity. Grasslands have more alkaline soils that

ENVIRONMENTS AND ECOLOGY OF MAMMALS 37 Table Env-2.Env-2 Some mammals and their soils, listed gen- encourage dense root development even when erally from north to south, and important beach dunes from dry; the result is called sod. In winter the entire Door County. prairie generally freezes and loses all green 1. Valders’ Clay and Loams including coarse northern soils and leaves and grasses, which must be renewed in some extending along Lake Michigan: Boreal species, such as the coming growing season. Herbivores and, snowshoe hare Lepus americanus, red-backed vole indirectly, insectivores or carnivores are affect- Clethrionomys, and star-nosed mole Condylura cristata dwell ed by this seasonal effect. there. 2. Northwest sands: pocket gopher Geomys, white-footed mouse Peromyscus leucopus COMMUNITIES 3. Peaty soils: Shrews, Condylura,Woodland jumping mouse Napaeozapus, southern bog lemming Synaptomys, muskrat Ondatra, and ermine Mustela erminea. Three Grand Floral Communities. In the simple binary subdivision of the Wisconsin envi- 4. Central Sands: P. leucopus, eastern cottontail Sylvilagus, prairie vole P. ochrogaster minor. ronments, the so-called “North Woods”, con- 5. Prairies, light loess about four feet deep more or less: Scalopus sisting of a boreal woodlands biome (decidu- aquaticus, Cryptotis parva, Reithrodontomys megalotis, ous, coniferous, and mixed forests) is north- Spermophilus, P. m.bairdii, Taxidea taxus. ward, and the prairies intermixed with decid- 6. Beach Dunes: P. maniculatus bairdii uous woods on the one hand, and on the oth- 7. Thin loess or forest topsoil of only a few inches depth where the er, many open agricultural fields and old fields, other soils are not exposed: Includes many mammals of prairie bottomlands, boreal remnants, local fens, and forest lands. bogs, and large marshes lie in the south. Open prairies are extensive grasslands; they not only look like great rippling seas, but are as adverse as seas to some mammals, while to others are essential. Woodland forms include eastern chipmunk, southern flying squirrel, and others. Prairie mammals include grazers, formerly including Bison, and numerous burrowers such as the prairie mole Scalopus, ground squirrels Spermophilus, and the prairie deer mouse, P. m. bairdii. Therefore, southern mammals seem divided as prairie and deciduous woodland species. Zimmermann (1991) in “Wisconsin Birdlife” discusses how three important floral areas meet in central Wisconsin: the boreal North Woods (mixed and conifer forest); in the southwest and south-central, the extensive dry prairies (intermixed with maple, basswood, prickly ash, and oak on the rugged highlands in the coulee country); and in the southeast part of the state extensive broad- leafed forests (beech, hardwoods). In Jones and Birney (1988) these same regions are termed: “hemlock-white pine-northern hard- Including Three Wisconsin Biomes (grand communities) woods” which in Wisconsin studies also have of North America. V. Shelford (U.Illinois Press). been called conifers and mixed woodlands; in

38 THE WILD MAMMALS OF WISCONSIN the southwest, known in Wisconsin more of- boreal kinds and recent southern invaders. ten as the Driftless Area or the Prairie-wood- There is much ecological diversity in forest lands, are many interconnected prairies and lands. Most northern communities are well extensive stands of “maple-basswood”; in represented by mammals, except Dry Mesic southeast Wisconsin are deciduous “Oak-hick- Forest and Dry Forest. The most depauper- ory forest” and “Beech-Maple” forest. ate is the Jack Pine Barrens. These have poor Curtis (1959) has written a classic study soils. Birch-Aspen Forest is not quite so well of plant communities of Wisconsin. Stearns represented with species as are the remain- and Kobriger (1975) extended this descrip- ing northern communities. The southern com- tive study for the watershed of Lake Mich- munities are not strongly represented by mam- igan. The analysis of the occurrence of mals, except Oak Savanna, followed by Mesic mammals in these plant communities fa- and Dry Mesic Forest. cilitates understanding of certain ecologi- There is much intermingling of habitats, cal relationships (Table Env-2). Some ex- even merging communities of the three ma- tirpated and abundant mammals are not jor biomes. For example, in central Wiscon- included in this survey. sin, where I have lived 40 years, there is a Lying along the Wisconsin moraine is an great deal of local plant diversity (spruce, hem- ecotone separating communities generally lock, white pine, sugar maple, red maple, but- into northern (N) and southern (S) groupings. ternut, pin and bur oak, hickory, cottonwoods, N1. Cat-tail or Reed Marsh // S1. Cat- aspens, bluestem as tall or taller than 2 m, tail or Reed Marsh basswood, paper birch, cat-tails, marsh grass- N2. Northern Sedge Meadow // S2. es, and jack pine in abundance). Likewise, in Southern Sedge Meadow central Wisconsin there is an amazing, large N3. Alder Swamp // S3. Shrub Carr (wil- mixture of mammal species (Long, 1970). low, dogwood) By use of average resemblance coeffi- N4. Northern Lowland Forest (tamarack cients to indicate faunal resemblance (of mam- and black spruce) and Relic Bogs // S4. mals), Long (1970) showed that in central Southern Lowland Forest (maples) Wisconsin the prairie assemblage of mammals N5. Northern Wet Mesic Forest (white resembles the Dry Forest edge (Oak Savan- cedar) // S5. Southern Wet-Mesic Forest na) most and the Jack Pine Barrens next. (American elm) These dry communities are not greatly dis- N6. Northern Mesic Forest (sugar maple) similar in terms of temperature, precipitation, // S6. Southern Mesic Forest (maples) floras and faunas. They are often combined. N7. Northern Dry Mesic Forest (white The community faunas seem to reflect gener- pine) // S7. Southern Dry Mesic Forest ally the faunas related to the aforementioned (red oak) physiographic provinces. N8. Northern Dry Forest (jack and red Tension Zone. In addition to describing pines) // S8. Southern Dry Forest (black oak) carefully and generally mapping the plant N9. Jack Pine Barrens // S9. Xeric Dry communities of Wisconsin, many of which are Prairie (sunflowers, bluestems) essential for the distribution of certain mam- N10. Birch-Aspen Forest // S10. Mesic mals, Curtis (1959) described an ecotone, Prairie (legumes, luxuriant grassland) which he called a “tension zone” extending N 11. Boreal Forest (spruce-fir) // S 11. across Wisconsin like a climatic belt. It is Oak Opening [savanna] (white and pin oak). roughly 50-60, as much as 100 miles (= 161 Nearly twice as many uncommon mam- km) wide. There is no clear explanation for mal species occur in the northern communi- this ecotone, except that boreal organisms ties as in southern ones. These include the extend southward to their limits and south-

ENVIRONMENTS AND ECOLOGY OF MAMMALS 39 ern organisms disperse northward into bore- with northern vegetation and southern ones al habitats of the north. Temperature plays a with southern vegetation. The vegetation is subtle role, and in earlier ages it may have related to precipitation and soil patterns. Ap- played a stricter determining role. The vege- parently there is some competition between tation types are correlated with the presence northern mammals and southern invaders or absence of animals, northern mammals coming face-to-face.

Table Env-3. Select animals related to plant communities. Revised Long 1974.

Species Plant community Comments Cryptotis parva S9-S11 Sandy loam, rare Peromyscus maniculatus bairdii ditto ditto Pitymys ochrogaster ditto ditto Taxidea taxus ditto + S2, N2, N9 ditto Condylura cristata N1-N6, N10-N11 Wet, mesic Sorex palustris ditto ditto, brooks, marsh Napaeozapus insignis ditto ditto Sorex cinereus ditto+S1-S5, S10 ditto Sorex hoyi N1-N11, S1-S8, S11 wet, dry, disturbed, water. Sorex arcticus N1-N4 Marsh, alders, Canary grass Blarina brevicauda ditto Most habitats Vulpes vulpes ditto ditto Myotis keenii, Wide-spread Needs hibernacula M. lucifugus, ditto ditto Eptesicus fuscus ditto ditto Lasionycteris wide-spread Forests, needs trichopterans, beetles, etc. Nycticeius trichopterans SE Wisconsin Ash, silver maple, oak Peromyscus leucopus ditto Woods, brush Clethrionomys N1-N6 Woodlands Microtus pennsylvanicus ditto Wet, mesic grasses Pitymys pinetorum 6, S6,S7 Leaf humus, clay soils usually Ondatra zibethicus ditto Wet, mesic Castor canadensis ditto ditto Sylvilagus ditto Brush Lepus americanus N1-N5, N11, ?N6 Wet, mesic, winter snow cover Canis latrans ditto + S1-S11 Ubiquitous Canis lupus N1-N11, formerly S1-S8, S11 remote Ursus americanus ditto Mast, remote Tamias s. doorsiensis Open woodlands Eutamias minimus N1-N6, N8-10 Open woodlands, rock outcrops, range retreating Marmota monax N6-N8, S6,S9, S11 Open woods, forest edge Spermophilus franklinii N1,N2,S1,S2, S10 Wet, mesic Lontra canadensis N1-N6, N10, N11 Wet, protection Lynx canadensis N1-N11 Forest, mesic, hares, winter snow cover Tamiasciurus N4-N11 Forest, savanna Glaucomys sabrinus N5-N10 Mast, tree cavities Synaptomys cooperi N6-N8, N11, S10, S1-S6 Grasses, sedges, black soils, boulders Erethizon N1-N11, S1-S8,S11 Northwoods, tree canopies Urocyon N4-N9, S4-S8,Sll Brush, rock outcrops Neovison vison ditto Wet, mesic Odocoileus ditto Most habitats Mustela frenata Usually N5-N8, S5-S8,S11 Mesic M. erminea N1-N5,?N11, S1,S3,S10 Wet, mesic, snow cover M. nivalis N1-N2, S10,S11 Rare

40 THE WILD MAMMALS OF WISCONSIN MICROHABITATS above or below the surface of snow. One can well imagine a diversity of microhabitats for all “Microhabitat” refers to one subdivision “pre- these kinds, which have one common protec- ferred” by a species above others within a larg- tion, a den or nest with a moderate microcli- er habitat. The word focuses on an organ- mate below ground and out of the wind. ism’s specific and localized environment, par- To conserve body energy, mammals may ticularly the physical parameters. Microhabi- use burrows (badger, moles, and pocket go- tat differs from “niche”, which is an organ- pher), lodges (beaver and muskrat), tunnels ism’s function in an ecosystem. under the snow (shrews, mice, and weasels), Not only have few microhabitats and their or caves (bats and occasionally other mammals) microclimates been studied in Wisconsin, even to provide more moderate environments. Even the nests of some obscure mammals have wolves and deer seek dense woods and shrubs, never been found or described. Heat and out of the wind, and lie partially buried with water availability play an unknown role for snow (Ozoga, 1968; Mech, 1970). Many mam- mammals in Wisconsin’s local habitats; but mals drag insulating materials into the den water metabolism is important in lactation, (bears, weasels, voles) or plug the openings to kidney function and heat loss. Summer provide additional insulation (badger, pocket drought and heat, inseparable problems, may gopher, ground squirrel), or dig down below become lethal and profound in the general the frost line (moles). Some huddle together to effect, but probably it is especially so, spottily conserve heat locally (e.g., pine voles, south- and sporadically important in variable micro- ern flying squirrel, the rare Indiana bat, and habitats. I have observed chipmunks basking occasionally striped skunks, raccoons, even in the morning sun on rocks that heat up to porcupines). Some mammals allow their go- 50o C on a summer day. nads to regress and cease breeding (deer mice As important as heat and water are, they Peromyscus, pocket gopher, some squirrels). are not so dangerous to mammals in Wiscon- At least three species of bats migrate south- sin as is the bitter winter. Starvation, disease, ward in winter (Lasiurus and Lasionycteris), and hypothermia lead to drastic mortality in while thousands of cave bats move into the exposed species during winter, and even for moderate microclimates of caves or opportu- those sheltered below ground if the frost is nistically into human buildings. deep. However, many mammals are well adapt- Few studies have addressed microclimate ed to winter and do not seem to suffer stress in homes of mammals. In a model study of the or high mortality. Some that do, seem to re- western ground squirrel (Spermophilus beld- cover. The white-footed mouse (Peromyscus ingi), Mohrhardt and Gates (1974) provide an leucopus) suffers terrible mortality in some example. Getz (1968) showed that microcli- winters in central Wisconsin, but it is the com- matic differences do not separate the red- monest mouse. Some mammals hibernate such backed vole and the white-footed mouse, but as ground squirrels (Spermophilus), wood- water use by the former causes it to prefer chucks (Marmota), and jumping mice (Zapus swamps compared to the drier uplands used and Napaeozapus); some cache food to last by the white-footed mouse. The microclimate through the winter such as the short-tailed has been studied in the winter lodges of bea- shrews (Blarina brevicauda), both chipmunks, ver (Novak, 1987) and muskrats (Huenecke, tree squirrels, the pocket gopher, beaver, and Erickson and Marshall, 1958), and in the dens some mice (Peromyscus); and some mammals of some large carnivores (Long and Killingley, forage below ground to eat roots and insects. 1983). Some studies of microclimates have Some mammals den up and slow their meta- been made in bat caves (Ainsley, 1983; Beer, bolic rates, whereas others forage actively 1955; Barbour and Davis, 1969). Promising

ENVIRONMENTS AND ECOLOGY OF MAMMALS 41 detailed, systematic and technological meth- (Long, 1965, and others). In Wisconsin, the odology is suggested by Bakken and Kunz microclimates are influential, especially in (1988). Whereas caves have been studied, the open forests. Owing to the layering of plant climate in the hollow trees or rock crevices used species, forests provide many more microhab- by bats has been seldom studied. The effect of itats and possible niches to specific mammals. the bat, itself, warming its immediate surround- This is one of the reasons species richness ings, and the warmth of the sun have been tends to be high in Wisconsin forestlands. seldom measured. The nature of the substrate Even the weather cycles of the ages, per- in the hibernaculum or maternity site of bats, haps from the long ago glaciations to this day, differing temperatures at various elevations, the may have evolutionarily fixed some respons- effects of wind currents and humidity, the es and other behavior, of hibernation and warmth possible from solar heat, heat from migration, to mention but two. We do not yet the bats themselves, depth of the cave and dis- know the role of long-term microclimatic fluc- tance from the entrance, and other factors all tuations on the Wisconsin mammals. divorce the interplay of variable microclimates We do not know the long-term interrela- from general weather. tionships of organisms within microhabitats, The temporal and spatial microhabitats the role of the millipede in the forest, of the theoretically provide opportunities for some sod worms and beetle grubs in the prairies, “lucky” or opportunistic mammals and disad- the role of shrews in the marshes. Lately some vantages for others: available foods of insects, conservationists, even the game managers of shoots, buds, seeds, fruits or mast; or home big game, are beginning to assess the small security of burrows, tree cavities, lodges, dead animals in the managed units. To understand falls, and dense shrubbery. Relationships also the total environment, the scheme of life, the include distance to standing water; relative importance of all members of our communi- humidity and snow fall; and open space space ties, and their relationship to humankind, it is to see and escape by running away or scurry- essential to study the cryptic and subtle ways ing up a tree for safety. All of these vary over of all the mammals and other life forms dwell- the landscape, and from season to season. ing in and relating to their microenvironments. One opportunistic example is provided by the Knowledge of microhabitats can be summed raccoon, known to use tree cavities and fall- up as a paucity. en logs for nests (Steuwer, 1943), but which adapts to prairies, savannas and marshes by Table Env-4. Ecological successional seres and selected building ground nests. Another example, the small mammals in them. Baker, 1983. white-tailed deer feeds opportunistically on plant droppings of a porcupine in winter (Sha- Mammal Crop Annual Perennial Shrubs Mature piro, 1949) or on plants thrown out of a Land Grasses Grasses Saplings Hard- Forbs Forbs wood muskrat’s opening on river ice (personal observations). The river current and condition Sorex cinereus ++++ of the stream banks may influence beavers to Blarina ++ + + + build lodges or to dig bank burrows; the same White-footed +++ dilemma affects nest building for the marsh- Mouse dwelling muskrat, which cannot tunnel where Prairie Deer ++ + the water table is high. mouse In the American West a mammal may be limited to a mountain, or even a northern Meadow Vole +++ slope of a mountain, or to a basin or desert Meadow Jumping ++++ by macroclimate, in a word temperature Mouse Zapus

42 THE WILD MAMMALS OF WISCONSIN ECOLOGICAL SUCCESSION ÀND LAND USE Species simply occupy favorable sites as they become available (Table Env-4-5). The Succession. Habitats change in sequential dynamic process of succession following stages called “seres”. These may take many ecological disturbance has permitted mammal years to elapse. Some mammals prefer Early populations to disperse into developing habi- stages, relatively brief, whereas others prefer tats here or there, while their own habitats the late (Climax) stages, which may last indef- changed adversely whence they came. initely (Table Env-4). Therefore, in good wild- Fire. Aridity and fire helped prairies de- life management the seral stages must be velop on many post-glacial soils, reducing maintained as well as the climax vegetation competition of grassland with developing for- (i.e., dominant vegetation that lasts). Early ests. Although prairie fires are formidable, and successional plants such as the transient black- I never forget the sight of rabbits and opos- berries, shrubs, aspens and seemingly insig- sums struggling to escape from one, fire prob- nificant understory and ground vegetation may ably does not destroy many of the mammals. be more essential to many mammals than the Mobile kinds may escape the flames and re- mature trees. Succession and its impact on turn. Some mammals hide in burrows, but the mammal habitats are illustrated (Table Env-4) prairie vole (Pitymys ochrogaster) may be for six common small mammals. adversely affected (although observed effects In Baker’s example, the shrew Blarina resulted from both fire and plowing). Pure was found in all five successional stages. Ex- prairie stands of grasses and forbs are cleansed cluding this shrew, only one mammal (Per- of encroaching thickets and trees, and the omyscus maniculatus bairdii) occurred in the prairie plants quickly return. earliest stage, and only three occur in the cli- In the northern forests, when the weath- max. If succession is staggered the fauna may er is dry, fire adversely affects many small include six mammals. mammals. In a three-year study following a

Table Env-5.Env-5 Choice of seral and climax communities in north (left) and south Wisconsin. Shade eradicates some mammals.

Mammal Shrub Cattail Sedge Dry Climax Sand Oak S. Dry Climax carr Meadow Pines Forest Barrens Barrens Prairie Forest

Sorex palustris XXX Lepus americanus XXX Sylvilagus fl. X XXX Marmota monax Rare X X X Rare Tamias striatus X XXXX Tamiasciurus XX X Sciurus niger XXXX S. carolinensis XXXX Glaucomys volans XX R. megalotis XXX Peromyscus m. bairdii XXX Peromyscus leucopus XXXXRareX Clethrionomys XX X Pitymys ochrogaster XX Meadow Vole X X X Rare Rare Rare X Mustela erminea XXX American Marten X X White-tail Deer XXXXRareXXRareX

ENVIRONMENTS AND ECOLOGY OF MAMMALS 43 huge forest fire (in the summer of 1976, of mammals in Wisconsin certainly benefit by the 60 square km) in the Seney National Wildlife appearance of grassy openings. The white- Refuge of Upper Michigan, Anderson (1982) tailed deer, eastern cottontail (Sylvilagus flori- showed an immediate adverse impact on nu- danus) and fisher (Martes pennanti) adapt merous small mammals. After the fire the to them especially well, and cottontails, fox most abundant small mammal was the masked squirrels and woodchucks thrive. White-foot- shrew (Sorex cinereus) and it was the only ed mice and southern flying squirrels may be species taken in the sedge marshes. By Octo- favored over their northern counterparts. ber 1977, the number of small mammals was Where fragmentation has created a mo- significantly higher in the burned habitats. saic of local habitats, the landscape is said to Meadow and red-backed voles were abun- be “patchy” and in scaling and complex pat- dant, as were chipmunks and other species. terns of food sources, cover, and distribution By 1980, an abundance of jumping mice of animals, these may exhibit fractal geogra- (Zapus hudsonius) raised the diversity even phy. Fractal analysis, a new approach to ap- higher. Through the succession, deer and praising any complexity that scales down and black bear seemed unaffected. The fire in- elaborates as it scales, may increase under- creased plant diversity, leading to an increase standing of such elaborate patterns. in small mammals. The savanna community is open grassy “Edge”, Patchy habitats, Savannas. In habitat with little continuous tree canopy. The ecology, the term “edge” has often been used scattered, mature trees are usually oak, in to describe organisms that dwell at the inter- some places jack pines, or various kinds of face of two distinct habitats. Usually the or- thickets. If the shrubby thickets are not con- ganism does not survive as well in either of tinuous, interfaces in the savannas may be the pure adjoining habitats. There are, of “edge” for numerous species of animals. course, many kinds of edge interfaces. In The openings made by humans (such as Wisconsin the most common is that between railroad or roadside rights-of-way, fields, lum- grassland and forest. The woodchuck Mar- ber camps) in forests may qualify as savan- mota is considered an edge species, because nas, and are often grown to weedy vegeta- it digs a den on an open slope providing vis- tion. Good examples of savanna mammals are ibility, easy digging, and access to plants for eastern cottontails Sylvilagus, ground squir- succulent foods. Forests may lack all three. rels Spermophilus, white-footed mice P. leu- Flat fields have poor drainage and less visibil- copus, the prairie vole Pitymys ochrogaster, ity if the vegetation is dense. The woodchuck and probably in western Wisconsin even the is often observed on grassy road embank- western populations of P. pinetorum. ments today, or on open hillsides with near- Forest Openings. Our northern forests by woods. Game managers recognize that are closing their grassy openings by ecologi- openings in the forest and old fields increase cal succession, especially on loamy soils. Thus, “edge habitats” which are favorable to deer succession causes deterioration of summer and some other game animals. Presently there range for many mammals. Usually 3-5 per- is used in scientific circles the trendy phase, cent of forest land maintained as open grassy which may last, “habitat fragmentation” or range would be of benefit to deer and other “landscape fragmentation”, suggesting or in- wildlife (McCaffery and Creed, 1969). Open- ferring that as “edge” increases, by whatever ings can be preserved easier than created. means, the pure habitat species are isolated Since the Wisconsinan glaciation, forest and often lost. This isolation needs much openings created by frequent natural forest fires study, for “clear cutting” and worn out fields have been extensive. Most of northern Wis- are becoming commonplace. However, many consin was forested by mature timber with lit-

44 THE WILD MAMMALS OF WISCONSIN tle undergrowth, but vast areas of barrens and northward in Wisconsin as far as Marinette burns were commonplace. Savanna-like shrub County, Michigan (Burt, 1948). Today that lands and pine barrens were widespread in the movement northward is accelerated, and northern forests. However, wet forests may some boreal forms such as the northern fly- show little effect of fire. On the Upper Penin- ing squirrel (Glaucomys sabrinus), the least sula a popular description is “asbestos forest.” chipmunk (Eutamias minimus), and the for- Today most forest openings are less than est deermouse (P. m. maniculatus) may suf- 10 acres in size and result from land use. In- fer because of competition. Water shrews and vading weedy species may thrive in such open- some jumping mice, and perhaps other mam- ings. Bracken grasslands and treeless expo- mals, may also be losing habitats due to land sures on sandy soil result from timber cutting practices and water pollution. Forest and wet- and fires. Many meadows on the heavier soils land preservation, if it continues in Wiscon- resulted from logging camps, pastures, and sin, may help save many of these mammals. failed farms. Plants present on loams include Excellent maps of former vegetation and the Agropyron, Poa, Cirsium, Rubus, and Achil- present day pattern of agricultural and urban lea. On medium soils today plants such as regions are given in a recent Wisconsin re- Poa, Hieracium, Agropyron, and Fragaria port (Anon. 1995, see maps). occur. On sand soils Hiracium, Myrica, Vac- Roadsides and railroad rights-of-way not cinium, and Poa are found. Other studies only are grassy openings in forest, but they show Pteridium, Myrica, Carex and Gault- provide linear (traversed faster) routes of dis- heria on bracken grasslands. persion. Many mammals (e.g., voles, deer These openings improve summer range mice, probably others) expand their range for white-tailed deer and provide habitat for a following these routes, and some (e.g., Mar- variety of small mammals such as voles and mota monax) build up their highest densities shrews, and carnivores preying on them. in such habitats (Woodward, 1990). The In the southern grasslands, and in the pine slaughter of mammals from mice to deer, in- barrens on thin soils are conspicuous assem- cluding opossums, raccoons, skunks and thir- blages of prairie plants. Notable are Andro- teen-lined ground squirrels, by automobiles is pogon, Liatris, Amorpha, Artemisia, Aster, excessive mortality. To manage rare mam- and Carex. Such plant communities allow free mals such as wolves along paved roads such dispersion of prairie mammals into the north- as Highway 53, in northwest Wisconsin, is a ern forests, and cause the strong affinities of primary cause of concern. prairie assemblages of mammals with those The Dairy Farm in Wisconsin, one of of the pine barrens and forest edge. America’s most unappreciated treasures, was Land Use. Land use such as clearing dense an almost biblical provision of wildlife habitat. forests, plowing fields and leaving some fields Dairy farms are found throughout Wisconsin. fallow, mowing rights-of-way along railroads A typical dairy farm may have woodland, open and roads, and lawns in urban areas, has al- fields and edge communities. The edge and lowed the recent invasion of many southern woodland provide wildlife over many years. species into the North Woods and up the Door Many farms failed, and their worn out soils lie Peninsula. By the middle of the 20th century, fallow. Prairie-like grasslands, thickets, and some southern species (e.g., Spermophilus forest may develop from the old-field habitats. tridecemlineatus, Glaucomys volans, Per- Lumbering nearly devastated Wisconsin’s omyscus leucopus, Peromyscus maniculatus northern forests early in the past century, and bairdii, and Sylvilagus floridanus) had found removed many of our finest trees throughout their way through open woodlands, wetlands the state. Clear cutting in strips today is a and along Lake Michigan dunes and beaches controversial practice that hurts some organ-

ENVIRONMENTS AND ECOLOGY OF MAMMALS 45 isms and benefits others; the harvest of old tive problems of raptors and DDT accumula- timber does the same thing. Replanting of the tion in their egg shells, which seemed to gal- forests and conservation by both private, state, vanize opposition to pesticides (1968-1969). and even county agencies have led to forest Frederick Baumgartner, George C. Becker, renewal and continuous use of some trees for Hickey, Hugh Iltis, Lorre Otto, Fred Ott, Leo- paper and lumber. Forestry was decidedly ni Vrtilk, William Reeder, and other support- beneficial to most Wisconsin forest and for- ers of the Citizens Natural Resources Associa- est-edge mammals. tion won this political battle. Urban mammals. In cities and suburban Other toxic substances that have been areas, besides the introduced Norway rats and studied in our ecosystems include PCB’s, house mice, and unwanted or free-ranging mercury, and numerous organophosphates. domestic cats, there are today numerous deer, These have both direct and indirect dangers raccoons, cottontails, big brown bats, chip- to wild mammals. For example, PCB’s poi- munks, cottontails, gray squirrels, voles, opos- son fishes, which are eaten by mink, raccoons, sums and occasionally other species that thrive and other mammals that feed extensively on in our midst. Besides the parks and boulevards, aquatic animals. even our finest estates tend to preserve some Bounty systems and other politics. natural habitats and native mammals to look Bounties have hurt some rather harmless spe- at, even in the largest cities. This coexistence cies. Today this waste of taxpayer’s money is of humans and wild mammals does not include curtailed. A valuable role of predators in any a great many kinds of mammals. Neither does ecosystem is now recognized. However, gov- it extend much to the mammals of the north, ernment often takes the side of businesses and the wetlands, vast prairies, or dense forests. urban development at a relentless expense of Mammals that fit well into the urban surround- nature, and there are more and more, wider ings, are those usually active and seldom seen and wider roadways. Some people are satis- at night that climb to safety into tree canopies fied if we save the big trees along the roads. or burrow in lawns, that have generalized di- But these old trees eventually will die. If most ets, perhaps including mast from desirable lawn wild mammals could choose saving the great trees, and den in areas of ecological distur- old elm or maple on the right-of-way or to bance. Some perceived as cute and beautiful save a patch of hazelnut, bluestem and cat- may become great nuisances. tail, they would not vote for the big tree. Why Insecticides and other contaminants. In not save some of the native shrubby, grassy this century natural history students must study habitat along roads, and focus wildlife toward the laboratory science of biochemistry, because roadside culverts? many problems of pollution and chemical con- Agencies involved with preservation of tamination of ecosystems will develop. Insecti- habitat include the Wisconsin Department of cides and other poisons such as DDT caused Natural Resources, state and private colleges great problems to wildlife, since their use after and universities, some enlightened county World War II, but their effects on mortality of governments, and several conservation soci- animal populations are not well known. Fortu- eties such as The Nature Conservancy. Envi- nately, the use of such poisons has been re- ronmental education is stressed by several stricted greatly. After Rachel Carson, in her natural history museums and nature centers. famous book Silent Spring, alerted the world In addition, the Wisconsin Academy of Sci- about the eminent danger of insecticides, Wis- ence, Arts and Letters Transactions was an consin became the first state to ban DDT. Jo- influential voice through Wisconsin history, seph J. Hickey and his student D. W. Ander- publishing many studies on Wisconsin’s natu- son studied at the UW-Madison the reproduc- ral history.

46 THE WILD MAMMALS OF WISCONSIN ZOOGEOGRAPHY some vanished from the warming lands. Lat- er, invasions of southern prairie and decidu- “So greatly has the climate of Europe changed that ous woodland floras alternated (Thomson, in Northern Italy, gigantic moraines left by old 1940). Frye et al. (1965) discussed glaciation glaciers are now clothed by the vine and maize... a in Wisconsin and Illinois. large part of the United States reveals a former cold With the new vegetation came the inva- period... As the cold came on, the temperate sion of new mammals from the south, a few inhabitants would be supplanted by Arctic produc- from the west, and a few that moved around tions. The inhabitants of the more temperate Lake Superior into the boreal mixed and co- regions would at the same time travel southward... niferous forests. Thus, a Holocene (i.e., Re- the present circumpolar inhabitants, which we cent) fauna rich in diversity was established. suppose everywhere travelled southward are Quite a number of interesting mammals be- remarkably uniform around the world... As the came extinct (Lundelius et al., 1983; Graham, warmth returned, the arctic forms would retreat 1976; Semken, 1983; see Prehistoric Mam- northward, closely followed up in their retreat by mals below). the productions of more temperate regions.” — The next upheaval of the mammalian fau- Charles Darwin, On the origin of Species, 1859. na came when Caucasians settled Wisconsin. Native Americans had perhaps helped to erad- Nowhere does Darwin’s profound and pi- icate some of the Pleistocene mammals, but oneer thoughts on glaciation and animal dis- insofar as is known, no habitats were destroyed tribution apply better than Wisconsin zooge- by them. White settlers, however, left hardly ography. The distribution maps compiled here- an acre of good soil unplowed, seldom a square in may be compared with those made by Jack- mile of heavy timber, or rarely a large game son (1961). The difference between the pat- animal or valuable furbearer remaining any- terns is mainly due to current effects of land where. Only in remote regions did these mam- use. That is the chief factor now encouraging mals linger, until people with conservation ide- some southern mammals to disperse north- als influenced society. That impact was pres- ward, and causing some kinds to vanish alto- ervation of forest reserves, reforestation, even gether. Zoogeography is a study of how the re-establishing prairies, and scientifically man- history and dynamics of climate and soils, both aging game animals. Another dramatic change past and present, affect the vegetation patterns, in mammal geography came in the past cen- which in turn influence animal distributions. tury from land use of modern society. Cities Since the history of mammal geography grow enormously, roadways multiply exponen- in Wisconsin begins in the late Pleistocene tially as does the population. Urban sprawl following the Wisconsin glaciation, there are occurs when people try to escape from the no records here of the early evolution of our cities, and many villages become towns, towns marsupial, shrews, rodents and so forth, which become cities, and everywhere people cut took place long ago and far away. Three dra- brush, plant lawns, and remove trees. DeVos matic phenomena subsequently have affect- (1964) discussed some changes in the known ed our mammals. First, amelioration of the ranges of mammals in this region. frigid Wisconsin glaciation, with subsequent Probably a general rule around the world, warming of temperature, outwash of glacial eradication of the largest meat mammals took meltwater, deposition of sands and loess, soil place early. Eventually some mammals were formation southward of sandy loams and eradicated for fur, or fear of their depredations. many infertile soils, and the establishment of Reintroduction follows that rule in reverse, the boreal forests chiefly spruce and fir. Some reintroduction favors the largest mammals (Ta- boreal mammals remained in the region and ble Zoog-1, bats excluded). However, land use

ENVIRONMENTS AND ECOLOGY OF MAMMALS 47 and climatic change have recently and adversely of central Wisconsin (Table Zoog-2). This map affected mammals of all sizes. showed two belts of grouped lines, one along Most state collecting of Wisconsin mam- the tension zone, the other a little southwest- mal specimens prior to the 1970’s was done ward. The more southern belt is basically a prai- around the UW-Madison campus. The U.S. rie boundary, for species (e.g., Bison) that did Biological Survey spent time and effort cir- not range as far north as the Tension Zone. cling the state to pick up all possible species In the second map, based on more re- that peripherally entered it. Although noting cent distributions discussed in this work, some some northward dispersion of mammals populations seem, unfortunately, to have van- through the Central Sands area, my students ished recently (Cryptotis parva, P. manicu- and I (1970, 1974) collected numerous bore- latus gracilis in the south, also Lepus al mammals and some southern kinds endemic townsendii, and Spilogale putorius). The there. This gives the impression that boreal major change primarily from land use in re- mammals unknown in the Central Sands, on cent years is a loss of species in the center— Islands of Green Bay, and elsewhere were of boreal (Sorex palustris, Eutamia mini- moving south. They were extensions of known mus, western glacial-sands dwelling Pitymys range, not dispersions of actual range. For ochrogaster minor), and southern affinity this reason, in my comparison with older zoo- (Reithrodontdomys megalotis, Pitymys pin- geography (prior to 1961, back to the turn of etorum, and perhaps others). Furthermore, the 20th century) I include my records show- numerous mammals recently (in historic times) ing the occurrence of mammals (1965-1970), have extended their actual ranges northward some of which now seem to be vanishing. (Didelphis virginiana, Sylvilagus floridanus, Semken (1988), even without knowledge Spermophilus tridecemlineatus, Sciurus of my records in the Museum Reports and else- niger, Glaucomys volans, Peromyscus leu- where, concluded that species richness in this copus, and others) and northeastward up to region is greatest near Curtis’ tension zone. The mixture of northern and southern, i.e., boreal versus prairie and deciduous woodland species, is fantastic in west-central and central Wisconsin. Such a rich fauna should become famous and prideworthy for Wisconsin. Jones and Birney (1988) wrote a chapter on zoogeography for mammals of the North- central states. They mapped the range limits (boundaries of known ranges) for over 90 species in a so-called “spaghetti” map of the region. They found the aggregation of boundary lines notably intense in three areas, including one in southwest and south-central Wisconsin. They too concluded that the mammal pattern was largely congruent with vegetation patterns, and the vegetation developed on interesting soils and features of the Post- Wisconsin glaciation. Spaghetti diagram showing range limits of Wisconsin I mapped my own “spaghetti” diagrams, mammals approximately 1900. The packing of species near one for Jackson’s (1961) maps with my early the tension zone is extraordinary for eastern North America. records included, especially based upon surveys See text.

48 THE WILD MAMMALS OF WISCONSIN the tip of the Door Peninsula and into Upper faunal areas extend well beyond Wisconsin’s Michigan (Didelphis virginiana, Spermophi- borders. Excluding the out-of-state species lus tridecemlineatus). used in their analysis, and many wide-rang- Bats are not credited much as records, ing species such as bats and mobile carnivores, because of their mobility and extensive wan- I also omit wide-ranging aquatic species such dering, but the southern forms Perimyotis as muskrats (Ondatra zibethicus) and bea- subflavus, Nycticeius humeralis, Myotis vers (Castor canadensis) (in its former range). sodalis, and even Lasiurus borealis are not common in the northern counties. Tree re- Table Zoog-1.Zoog-1 Body size and extinction or eradication moval in towns and along boulevards may of known animals. Pleistocene to Recent. Game management have somewhat restricted the numbers and saved some species. Only game animals and furbearers have range of the red bat, even in the south, for been reintroduced. Mammals, weights, times. Jackson (1961) reported it was the most abundant bat in [southern] Wisconsin. Jones and Wooly mammoth 5,000kg, Late Pleistocene Mastodon 5,000 kg, Late Pleistocene Birney (1988) used faunal “units” for assem- ?Ground sloth 3,000-5,000kg, Late Pleistocene blages relating regions to their mammals. A Pleistocene bison 908 kg Pleistocene region is considered the origin for the mam- Plains bison 386 kg Late Pleistocene-historic mal in our state if the mapped geographic American elk 363 kg Late Pleistocene,Historic range extends (peripherally) into this state Black bear 272 kg Late Pleistocene-Present Pleistocene caribou 227 kg Late Pleistocene from that direction. Widespread species have Woodland caribou 182 kg ?Historic a more dubious origin. Ecological require- White-tailed deer 113 kg Late Pleist.-Present ments, such as strict preference for grasslands, Cougar (Mountain lion) 91 kg Historic were used to help form these assemblages Giant beaver 91 kg Late Pleistocene (i.e., “faunal units”). In their study of seven Timber wolf 55 kg Historic, reintroduced American beaver Historic, vanished in south states ranging from the Canadian border to Canada lynx 20 kg Historic-Present rare southern Illinois and Indiana, some of their Wolverine 18 kg Late Pleist. –?Historic Coyote 18 kg Historic-Present Bobcat 18 kg Historic-Present Raccoon 18 kg Historic-Present American badger 11 kg Late Pleist.-Present Fisher 6.8 kg Historic, Reintroduced Red Fox 6.8 kg Historic -Present River otter 11.3 kg Historic – Present Striped skunk 4.5 kg Historic-Present White-tailed jack rabbit 5.4 kg Historic-Present American marten 1.4 kg Historic, reintroduced Mink 1.5 kg Historic – Present Muskrat 1.5 kg Historic-Present Large weasels 100, 200 g Historic-Present Eastern chipmunk 100 g Historic-Present Meadow vole 50 g Historic-Present Vole P. pinetorum 40 g Historic-Present Vole P. ochrogaster 40 g Historic-Present Phenacomys 40 g Pleistocene Peromyscus 24 g Historic-Present Blarina shrew 18-24 g Historic-Present Water shrew 12-18 g Rare Historic-Present Harvest mouse 12-16 g Historic-Present Masked shrew 4-6 g Late Pleist.-Present Spaghetti diagram for species from approximately 1960, Pygmy shrew 4-5 g Historic-Present rare showing range changes. Cryptotis shrew 5 g Historic ?eradicated

ENVIRONMENTS AND ECOLOGY OF MAMMALS 49 1. Widespread mammals — most of North tis parva and Scalopus aquaticus; bats Per- America and having obscure direction of origin. imyotis subflavus; Nycticeius humeralis; 2. Eastern widespread — deciduous for- cottontail Sylvilagus floridanus; and rodents ests of the southeastern United States. Tamias striatus; Marmota monax; Sciurus 3. Eastern Austral — extreme southeastern carolinensis (but perhaps it invaded from United States. Perhaps Nycticeius humeralis. New England?); Sciurus niger; Glaucomys 4. Eastern New England — Northeast volans; Peromyscus leucopus; and Pitymys United States and southeastern Canada. pinetorum. 5. Plains/Grasslands — prairie forms Some of the above species now have from the south and west. occupied the entire state of Wisconsin and 6. Boreo-montane — Forests of the Rock- much of the Upper Peninsula. Most of them ies or Canada. were, because of Lake Michigan interven- Widespread Mammals. In Wisconsin, the ing, from southern invasions, and are ap- widespread mammals of obscure origin in- proaching or invading the boreal forests of clude bats Myotis lucifugus, Lasionycteris the North Woods. Cryptotis and both Pity- noctivagans, Eptesicus fuscus, Lasiurus mys, however, seem eradicated or at least borealis, and L. cinereus (which seems a in peril. For the most part, land use favors southern species to me), wide ranging semi- these eastern species. This faunal unit aquatics Ondatra zibethicus and Castor ca- thrives on roadways, park lands, and forest nadensis, deer mouse Peromyscus manicu- edge in modern Wisconsin. latus (which acts as a species pair, i.e., both Eastern New England faunal unit. Ap- a boreal and a southern species in Wiscon- parently this assemblage had origin in the sin), porcupine Erethizon dorsatum, and vicinity of the Gaspe Peninsula, and perhaps carnivores Canis latrans (which seems south- they even came from the north. Those spe- ern in Wisconsin in relation to the wolf), C. cies occurring in Wisconsin or approaching lupus, Vulpes vulpes, Urocyon cinereoar- from northeast Minnesota have circled genteus, Ursus americanus, Procyon lotor, around the north shore of Lake Superior and Mustela frenata, N. vison, and Taxidea tax- invaded from the west. Therefore, to say they us (which is southern, perhaps Neo-tropical are eastern is like saying black is white; but in origin except it was ages ago adapted to their origin is east. These include Blarina this frigid land even in the Pleistocene Ep- brevicauda; Condylura cristata; Myotis och). Mephitis mephitis, Lontra canaden- keenii; Synaptomys cooperi; and Napae- sis (which is a northern Wisconsin species ozapus insignis. today), Felis concolor (which was eradicat- With the possible exception of the shrew ed), Lynx rufus (which is more southern pos- Blarina, all these mammals are adapted to sibly because of the boreal Canada lynx, but the northern conifers and mixed forests. today has moved northward), Cervus elaphus They resemble in habitat the Boreo-mon- (eradicated), and Odocoileus virginianus. tane element. There probably would be no objection to Plains/Grassland faunal unit. These in- consider Peromyscus maniculatus bairdii clude: Lepus townsendii (probably eradicat- and Lasiurus borealis as southern species, ed recently); Spermophilus franklinii (south- the former as a Plains/Grassland form, the ern grassland); Spermophilus tridecemlinea- latter perhaps the same (but it is highly mo- tus (southern grassland); Geomys bursarius bile, always associated with trees, and migrates (western); Reithrodontomys megalotis far to the southward). (southern or western); Pitymys ochrogaster Eastern widespread faunal unit. East- (one race is western, the other southern); ern species include the insectivores Crypto- Spilogale putorius (probably eradicated,

50 THE WILD MAMMALS OF WISCONSIN western); Bison bison (eradicated, western) from refugia in southern and eastern (what is (see section on Eradicated Mammals). now) United States. Paleontological evidence Jones and Birney (1988) listed Reithrod- proves the species has been here in early ontomys megalotis from the Southwestern Wisconsin boreal habitats. That occurrence Faunal Element, because the harvest mouse supports Stewart and Baker’s statement, ranges into Mexico. It is a grasslands species based on the DNA “clock”, that the differen- that invaded Wisconsin arriving from south- tiation of these Sorex populations was prob- ern or western states. Not discovered in Illi- ably before early Wisconsin time. nois until the 1950’s, the harvest mouse has Origin from the Deep South. The adapt- moved eastward and northward. From south- able opossum (Didelphis virginiana) arrived ern and southwest Wisconsin it dispersed into centuries ago from a South American ori- central Wisconsin. It is restricted to prairies, gin. It has invaded the northern counties, meadows, old fields, hayfields, and marshes. and finally reached Lake Superior, in up- The mouse seems to have a western-south- per Michigan. western affinity with Geomys, Spilogale and Zoogeography of Wisconsin Islands. the western race of Pitymys pinetorum, all The zoogeography of the islands at the from the Great Plains. mouth of Green Bay was discussed by Long Boreo-montane faunal unit. Even with- (1978). New species of mammals (endemic out including the somewhat boreal Eastern Synaptomys cooperi jesseni, introduced New England assemblage, this boreal and Urocyon cinereoargenteus, and Peromy- montane assemblage is the majority of Wis- scus leucopus) are recognized from Wash- consin mammals: Sorex arcticus; Sorex ci- ington Island. Unlike the Beaver Islands, nereus; Sorex palustris; Sorex (Microsor- more central in Lake Michigan, this group ex) hoyi; Lepus americanus; Eutamias min- called the Grand Traverse Isles had a few imus; Tamiasciurus hudsonicus; Glau- endemic kinds in the early Holocene (i.e., comys sabrinus; Clethrionomys gapperi; Recent). They arrived by Fox River rafting, Microtis pennsylvanicus; Zapus hudsonius; swimming, and crossing of the Lake ice in Martes americana; Martes pennanti; Mus- winter. Bats fly freely from isle to isle. Most tela erminea; Mustela nivalis; Lynx ca- hibernators are absent. Some kinds were nadensis; Alces alces. introduced by humans (Long, 1978). The two Martes are now reintroduced. Long (1978) reported that: 1. Only a few The woodland caribou Rangifer tarandus and species were present on most islands, except wolverine Gulo gulo have possibly historic the large Washington Island. 2. Often a spe- records in Wisconsin; they also fit in the bo- cies was exceptionally abundant. Never were real faunal assemblange. Boreal Phenacomys two species abundant at the same time, ex- intermedius might range into northwest Wis- cept very different kinds such as red-backed consin from adjacent Minnesota. Although voles and little brown bats on Poverty Island Sorex cinereus is obviously of boreal origin, or deer and raccoons on Chambers Island. 3. and restricted from some southern habitats The abundant species often had a widened in Wisconsin, the evidence from mitochon- ecological niche. For example, red-backed drial DNA (Stewart and Baker, 1997) suggests voles on Big Summer Island, Michigan, tun- Michigan shrews, and presumably Wisconsin neled extensively. 4. Some species occurrenc- shrews, show intermediacy between western es were unexpected, probably from chance populations (including Sorex haydeni ) and rafting. Condylura on Big Summer Island is those from eastern Canada (and New En- a good example. 5. Introductions by man was gland). The masked shrew probably invaded often a factor (e.g, Mephitis mephitis). 6. Canada after the Wisconsinan ice receded, Animals that crossed lake-ice occur on most

ENVIRONMENTS AND ECOLOGY OF MAMMALS 51 if not all islands. 7. Large islands with diverse jesseni, and Peromyscus maniculatus man- habitats have the most species present. 8. Is- iculatus (= gracilis). The forest deer mouse, lands in Green Bay resemble the Beaver Is- driven off the Door Peninsula by P. leucopus lands, eastward in Lake Michigan, only in the (Long, 1996), persists on most of the isles boreal mammals present. 9. Boreal mammals (Long, 1974a; 1978 c; 1978a; 1990; 1996; probably colonized these islands by crossing Long and Long, 1993). ice and by rafting, not by land bridges con- Condylura cristata probably arrived at necting isles together. However, Rock and nearby Big Summer Island from Upper Mich- Washington islands were once connected, and igan. Numerous mammals swim or cross the show faunal resemblance. On Washington Is- ice to these isles from the north shores of land are 15 species (including new P. leuco- Green Bay, mostly Upper Michigan (Ursus, pus, Urocyon cinereoargenteus, and Syn- Lynx rufus, Urocyon, Vulpes, Canis latrans, aptomys cooperi), on Rock 8 species (new Lepus americanus, and others). The bog lem- S. cooperi), and nearby in Michigan waters ming, Synapatomys cooperi, probably was St. Martens Island (5), Poverty Island (4), Big endemic for thousands of years because it Summer Island (8), and Little Summer Island shows some speciation. (3). Chambers Island, in Green Bay, had six. The greatest faunal resemblance to the The Garden Peninsula of Michigan had 23 Peninsula fauna, using the simple ratio of species, and the north end of the Door Pen- number in common to total compared, is that insula had 29, and they had a strong faunal of Washington Island (50 percent) and the resemblance. least is that of tiny Poverty Isle (only 4 per- Resemblances were determined by a for- cent). Again we find that Rock shows a high mula (Long, 1963) allowing comparison of affinity to Washington Island. All its kinds large and small assemblages fairly, without so occur also on Washington Island. Comparing much influence from the large-value denomi- the fauna of the island group to that of the nator due to the larger faunal assemblage. The Faunal Units of Jones and Birney’s definition, first percentage is that of the Peninsula assem- all bats excluded, the results are given below. blage, the second is of the Wisconsin assem- Boreo-montane 18%/ 36%; New En- blage as listed by Jones and Birney (bats ex- gland 7*/75*; Eastern 6**/22***; Plains/ cluded). The percent resemblances are in Ta- grasslands 0/0;Wide-ranging 25/30. ble Zoog-3. High values were found for Rock, These percentage values are influenced Washington, and St. Martin’s islands. Big Sum- by low numbers of Southeast, Grasslands and mer and St. Martin’s islands had strong faunal Neo-tropical units. The denominator is small, resemblance. Across Green Bay, the boreal so the percentage is high. Garden and Delta peninsulas compare closely The north end of Door, decidedly less with the boreal north end of the Door Penin- prairie-like than at the base of the Peninsula, sula (containing Erethizon, Clethrionomys, has these affinities with Wisconsin units (bats Glaucomys sabrinus, and others). excluded): Three island species are unknown on the Eastern 21% ; Eastern New England 25 north end of the Door Peninsula, surprising- % (only a single kind, Blarina); Plains/grass- ly, because it is so close to nearby Detroit, lands 4 %; Boreo-montane 25% . Washington, and Chambers islands. These are Much like mammals, not mobile and inac- Condylura cristata, Synaptomys cooperi tive in winter, the “herps” (reptiles and am-

* The bat Myotis keenii would swell this resemblance if bats were included. ** 7 percent if the recent white-footed mouse was counted. *** Includes the gray squirrel, which seems to me a New England kind.

52 THE WILD MAMMALS OF WISCONSIN phibians) of this region faced similar difficul- mals receding northwards will vanish in nu- ties in colonizing islands. Long and Long (1976) merous counties within their former range. and Long (1994) studied the zoogeography of The forests in the north are now favorable herps of the islands in Lake Michigan, finding to these species. little affinity with western Wisconsin herps, The interesting and diminutive glacial- quite a strong resemblance to those from the sands race of the prairie vole, Pitymys ochro- south, and some to those from the east. The gaster minor, has been greatly decimated. fauna of the land of Door, both peninsula and Its status is confused, probably extant, but the isles, is a mixture of boreo-montane and de- whereabouts at present are unknown. A few ciduous woodland mammals or herps. other species have vanished or become rare. Surveys in the Apostle Islands have proved Most of the Eastern Widespread species are less interesting. Little speciation is apparent doing well and will likely continue to do so. and the small mammals and most bats are bo- The introduced mammals may continue real. The red bat may not be long established. to flourish, except Lepus townsendii. Mar- Only a few species of mammals are present. Zoogeography Of The Central Sands. Comparing the rich tension zone fauna of the Table Zoog-2. Some species used for spaghetti diagrams Central Sands area, within the square area of and faunal estimates. Wide-ranging species excluded. Origin Marathon, Wood and Portage counties, with (N, S, W) and species. the list in table Zoog-2, the faunal resemblance Opossum N (bats excluded and extirpations not considered) Star-nosed mole N or NE is 91 percent (N 33). That is extremely high, Prairie mole S, SE a rich mixture of all kinds of mammals, as Arctic shrew N mentioned above. Central Wisconsin mam- Water shrew N local, rare Pygmy shrew N Uncommon mals of aforementioned Faunal Units [percent Least shrew S, SE extirpated of central species/percent of faunal unit] are: White-tailed jack rabbit W eradicated Neotropical 1, 3%/50%; Plains/grasslands 4, Snowshoe hare N 12/66; Eastern widespread 9, 27/90; East- 13-lined ground squirrel S ern New England 4, 12/100; Boreo-montane Franklin’s ground squirrel S, scarce Least chipmunk N 17 50/100. Red squirrel N This species-rich savanna area with bo- Fox squirrel S real forest northward and hickory-maple, oak- Southrn flying squirrel S, SE jack pine, wet and dry grasslands, and other Northern flying squirrel N habitats, has a fauna involved with all the Plains pocket gopher W Harvest mouse W, SW uncommon aforementioned units. It is primarily eastern Prairie deer mouse S (deciduous woodlands) and boreo-montane. Forest deer mouse N, NE Some grasslands forms are present, and the White-f ooted mouse S, SE opossum from the southern United States is Pine vole S, W also. Rare now well established. Prairie vole S, W also. One race rare. Red-backed vole N Future Changes And Grounds For Op- Woodland Jumping mouse NE timism. The future biogeography of Wiscon- Porcupine ?N sin will probably show a proportion of the Ermine N mammal fauna invading farther north, some Least weasel N Scarce reintroduced carnivores (and also the moose) Spotted skunk W River otter N dispersing farther south, and other mammals Canada lynx rare NW driven northwards by the advancing south- Bobcat S ern mammals. Some boreo-montane mam- Moose N, NW

ENVIRONMENTS AND ECOLOGY OF MAMMALS 53 tes foina may expand its range and numbers. vive. In Door County, southern invaders will There need be no protection provided for move up the peninsula, and some may cross Mus, Rattus, or feral cats, all of which should to nearby isles. Some species there may not be humanely disposed. survive (Erethizon dorsatum, Glaucomys Along the southern border of Wisconsin, sabrinus, several carnivores) because of land we may expect rare bats to retreat southward, use and growing population there. even if the temperature continues warming. The rich species diversity in central Wis- They are incompatible with intense urbaniza- consin may lose some species. Who cares what tion and sprawl. Some western species may happens in the center? With future efforts of extend their ranges eastward across Wiscon- our modern Wisconsin natural resources agen- sin, crossing and working northeast along the cy, I expect that few species will disappear from rivers, if they can cope with land use. One or Wisconsin. Probably some eradicated mammals two of the “New England” kinds may enter may be transplanted and reintroduced. There Wisconsin in the northwest, from Minnesota. are so many lovers of nature in this wonderful In the forested north most kinds should sur- state that one can guarantee a happy future.

Wooly Mammoth. Woodcut G. Cuvier. Regne animale.

54 THE WILD MAMMALS OF WISCONSIN PREHISTORIC MAMMALS with the pine and oak forests, especially throughout the southern half of present Wis- All of the prehistoric mammal kinds of what consin. Open country became prairie lands is now called Wisconsin invaded from outside and wet marshes. Forests on Valders moraines this region. Their Wisconsin history does not joined those of Cary moraines. The Door Pen- extend far back in geological time, their fos- insula and the surrounding waters were un- sils are unknown in our long-buried sedimen- covered of ice. tary rocks. In many places the heavy glacial Evidently early humans arrived about this ice of ages past scoured away these rocks, time. Since then the climate has fluctuated, be- and the resulting outwash sands buried other coming even warmer than we enjoy at present. rocky strata. The early history of Wisconsin Giant beavers (Castoroides), which did mammals must be traced back through time not build dams or lodges, lived in glacial melt- in far away places. water lakes (Dallman, 1969). Two huge mem- The Pleistocene glaciations, mentioned bers of the elephant family, the plant-forag- earlier in the Environment section, were, in ing mastodon (Mammut) and the woolly mam- chronological order, the Nebraskan, Kansan, moth (Elephas), ranged through the wood- Illinoian, and finally, and least extensive of the lands and dense spruce forests near the ice four, the Wisconsinan. Many Kansan and Ne- fronts. The mammoth probably ate grasses, braskan deposits are confused and often iden- leaves, and twigs. Giant flat-headed peccar- tified as “Pre-Illinoian.” The last stage, the ies (Platygonus ), huge bison (Bison occiden- Wisconsinan glaciation, probably ameliorated talis), a large boreal caribou (Rangifer (melted, retreated) in southern Wisconsin some tarandus), and even musk oxen (both Sym- 15,000 years ago, and the complete withdraw- bos and Ovibos) inhabited the boreal wood- al of ice from northern Wisconsin and the Great lands near the retreating ice fronts. The lem- Lakes occurred about 10,000 years ago. This ming Dicrostonyx, a small rodent, lived in marked the end of the Pleistocene, and com- this region, and was preserved at many Pleis- menced what is called Recent time (also called tocene sites outside Wisconsin. Now it is con- the Holocene Epoch). fined to the Arctic. One huge mammal not There were several advances of Wiscon- yet found or reported in Wisconsin is the gi- sin ice. Pollen analysis and carbon dating have ant ground sloth (Megalonyx). The elk-like helped us understand Holocene events by the long-legged moose (Cervalces scottii ) was occurrence of certain trees and grasses. Near recently found in Marathon County, buried Green Bay (Two Rivers, Peter’s Quarry, Duck where the Wisconsin ice front had melted Creek) and westward near Iola (the Iola Bog) away (Long and Yahnke, in press). Several bogs provide a record of former plants and species of whales and the walrus swam in Lake pollen. It shows that black spruce, tamarack, Michigan (Handley, 1953) and probably visit- and some grasses (Cyperaceae) were replaced ed the western shore. [I examined a Pleis- 11,640-11,850 years BP [BP= before the tocene whale vertebra from a Wisconsin gravel present], after the Cary ice advance and dur- pit, but the antiques dealer who owned it re- ing the warmer Two Rivers Interval by a shift fused to divulge the locality of this “valuable to more composites and dry prairie grasses. dinosaur find.”] After the last advance of ice, called the Valders Both warming climate and activities of Re-advance, the great invasion of the region early human cultures seemed adverse to the by white pine and oak (Schweger, 1969) com- large (and probably shaggy) ice age mammals, menced. Boreal forests and probably boreal for many became extinct. They left only teeth mammals gave way to, or were mixed with, and bones buried in bogs, swamps, and flood the arrivals of southern mammals associated plain soils to document their presence. Many

PREHISTORIC MAMMALS 55 medium-sized and small species persisted, The Moscow Fissure in the “Driftless some retreating northward following the re- Area” yielded fossils carbon-dated at roughly treating cool conditions. Others from the 17,000 BP, at the peak of Wisconsinan gla- southern hardwood forests (deciduous forest ciation (Rassmussen, . The hilly area was prac- species) and steppe species (Hoffmann and tically encircled by massive ice fronts, both Jones, 1970) from the vast interior grasslands westward and eastward. Many boreal and (or Great Plains) invaded the “uncovered” and Arctic species and genera were present, in- open habitats. Some deciduous trees and cluding Dicrostonyx, Synaptomys, and mammals may have overextended northward Phenacomys, including some species known in the warmest seasons, as oaks, hickories and today far to the westward (Table Pre-1). There other southern plants established themselves are no records of Deciduous-Woodland spe- into northern Wisconsin, some mammal pop- cies. The number of species of shrews and ulations persist today as relicts. voles is high, suggesting boreal climate (Gra- Most boreal species in Wisconsin today ham 1976). The Moscow Fissure fauna inhabit northern spruce, pine, maple, and showed greatest affinity with mammal com- eastern beech-maple forests. There is a rich munities of present-day, southeast Manitoba “amalgam” of these and warmth-adapted spe- and southwest Ontario (and perhaps north- cies especially in southwest and central Wis- ern Minnesota). consin. Along Wisconsinan ice fronts the From one site in western Wisconsin, Holocene and late Pleistocene fossil sites re- now destroyed to make a park, Palmer veal diversity (Semken 1988; Thompson, (1954) discussed 300 Bison occidentalis 1940; Foley, 1984; Thieling, 1973; Rasmus- that were associated with Native American sen, 1971). Whether microhabitats were di- remains. He (1974) also found the giant verse, free intermingling of the post-Pleis- peccary Platygonus. tocene mammals, or climatic fluctuation The fauna of Lost River Sink (Table Pre-2) caused this species stacking is uncertain. seems similar to the same fauna found there Some small fossil assemblages were con- today. At Lost River Sink, at level 3, the age is centrated together (by means unknown, prob- 2,720 BP, and at level 4, it is 3,970 BP. The ably by snakes, in some cases by owls, possibly by ants) in rock fissures, sinks, and caves. Fragmentary parts of the skull and teeth are Table Prehist.-1Prehist.-1. Recent and former species’ facies from most abundant at such sites. Cave deposits the Moscow Fissure, Wisconsin. This area not covered with on the Door Peninsula carbon-dated at one glacial ice. However, the fauna changed drastically. All the level at 600 BP, according to Robert Howe older species either retreated out of Wisconsin, or live mostly (personal comm.), contained numerous rein northern counties. Sorex hoyi ranges south. mains of some modern-day mammals, prob- 17,000 BP Recent species ably even Pitymys pinetorum. Mastodons excavated by Dallman (1968) Sorex arcticus Didelphis virginiana Sorex palustris Cryptotis parva near Deerfield, Wisconsin, in Dane County, Sorex hoyi Scalopus aquaticus were aged as 9,480 BP, and 10,095 (or a lit- Thomomys talpoides* Tamias striatus tle older). Dallman aged a woolly mammoth, Clethrionomys gapperi Glaucomys volans roughly contemporary in time and place with Synaptomys borealis* Geomys bursarius the mastodons, as 9,065 BP. This elephant Phenacomys intermedius Reithrodontomys megalotis Microtus xanthognathus* Peromyscus leucopus was found at Lake Mills, in Jefferson County. Dicrostonyx torquatus* Pitymys pinetorum Other large mammals (Bison occidentalis, Zapus princeps* Zapus hudsonius Castoroides ohioensis, and Rangifer tarandus) are likely as old. *No longer in Wisconsin

56 THE WILD MAMMALS OF WISCONSIN Table Prehist-2Prehist-2. The Lost River (L) , Moscow Fissure (M), and other sites are represented by numerous species. The older species from M are shown here. L seems recent.

Sorex arcticus M, S. palustris M, S. hoyi M, S. cinereus M, L, Myotis sp. L, Perimyotis L, Lepus americanus ?M, Thomomys talpoides M, Sciurus sp., L, Tamias striatus L, Spermophilus tridecemlineatus L, Tamiasciurus hudsonicus M, Peromyscus maniculatus L, Synaptomys cooperi L, Clethrionomys gapperi M, Pitymy pinetorum L, P. ochrogaster (large race) L, Phenacomys intermedius M, P. ungava (M?), Castor canadensis, Castoroides ohioensis, Mammut americanun, Elephas primigenius, Tayassu, Platygonus compresses, Cervus elaphus, Rangifer tarandus, Bison occidentalis (= latifrons), B. bison, Ovibos, small mammal fauna, then, has been fairly sta- Symbos cavifrons, ble for nearly 4,000 years in southern Wis- Ursus americanus L, consin. The modern species are, of course, Procyon lotor L, described in their species accounts beyond. Elk Mephitis mephitis L, Vulpes vulpes L, antlers from Portage County were buried after Canis cf. lupus, being shed along streams, and may not be of Gulo gulo, Jackson 1961. great age. A huge subfossil specimen from NE Columbia County is in the UW Wildlife Ecolo- gy collection. I saw a bison tooth from a field near the Wolf River east of Stevens Point, but there is no certain evidence of a natural burial. Archeological digs often yield some fossil remains, and many fossils are discovered by accident (in excavations). Attention to cave deposits and fissures provides many Recent fossils and subfossils. The small mammals are often obtained by washing and screening techniques. The huge bones of the large species are often found buried in lake beds, bogs, and Pleistocene Bison latifrons. After Hamilton, 1939.

PREHISTORIC MAMMALS 57 covered it should be immediately reported to Bison bison. William T. Hornaday. His writings probably professional geologists, and if possible left saved this species. It is now domesticated in Wisconsin. undisturbed. Fossils collected without accompanying data are worthless. Usually a plan can flood plain sediments where the smaller forms be worked out to accommodate all interested are not preserved. When a fossil site is dis- parties at a fossil site.

Pleistocene and post-Pleistocene records of large Ice Age mammals showing Mammut mastodon; Elephas mammoth; Bison; and Rangifer caribou. The Wisconsin drift and lobes are also shown. The Bevent site contained two caribou and the only Cervalces scotti elk-moose (above) known in Wisconsin. The fossil elephant sites lie south of the Laurentide Wisconsin ice sheet, or near it as it receded. The large caribou are from the glaciated southeast or central part of what is Wisconsin, found in wetlands near moraines. Bison, as in pioneer times, ranged along the west border of Wisconsin as this region was to become prairie and woodland. West and Dallman, 1980, Long, 1986, West, 1978. Map and photo by C. Yahnke.

58 THE WILD MAMMALS OF WISCONSIN REINTRODUCED AND EXTIRPATED The reintroduction of small mammals has not MAMMALS OF WISCONSIN WITH yet begun (Long, 2001). HYPOTHETICAL LIST OF SPECIES Some mammals introduced or reintro- POSSIBLY IN WISCONSIN duced in Wisconsin are surviving, and even thriving. They are mentioned here and treat- How does one know whether secretive speed in detail in the species accounts below. cies are unknown or eradicated? Some spe- Some eradicated native kinds reintroduced are cies are very difficult to detect. This listing sometimes of a different geographic race (e.g, supplements the list of earlier Pleistocene and Cervus elaphus nelsoni). If eradicated before early Post-Pleistocene mammals eradicated by historic time, the species are discussed in Pre- warming climate, human activities, and un- historic Mammals. If eradicated in historic known causes. Eradication of mammals from times, up to approximately 1900, they are Pleistocene and Holocene Wisconsin gener- included immediately below. ally was related directly to the mammal’s body Cryptotis parva. Least shrew. This shrew mass (probably important both for cold weath- may have vanished about 1940. See account er subsistence and for the Native Americans of the least shrew. for food—Table Zoog-1), climatic warming Lepus townsendii. White-tailed jack rab- (which affected Ice-Age mammals regardless bit. Possibly native in southwest and north- of their mass), fur use, intolerance to preda- west Wisconsin, introduced in many places tors by farmers and hunters, and land use (i.e., in the state, and possibly eradicated about habitat alteration, which is affecting the hold- 1970. See account of this species. overs from earlier eradications). Natural and Rattus norvegicus. Norway rat. An in- scientifically managed reintroduction gener- troduced and established pest. See account ally follow the same rules (without any rein- of this species. troductions of Arctic species, of course), gen- Mus musculus. House mouse. An intro- erally in the same sequence, but eradication duced and established pest. See account of of the small mammals may be far from end- this species. ing. Their geographic ranges are constantly Martes americana. Pine (or American) Mar- changing, favoring in some cases the species ten. See account of this reintroduced species. scientifically managed, with some others in- Martes pennanti. Fisher. See account of directly benefiting from the management, this reintroduced species. many southern invading species, such as Glau- Gulo gulo. Wolverine. Possibly eradicat- comys volans, and some that adapt well to ed. Jackson (1961) reviewed the history of modern land use (e.g., Didelphis, Eptesicus, this problematic, large mustelid, under the Sylvilagus, Spermophilus, Sciurus niger). name Gulo luscus luscus, the only specimens of which are sub-fossils from cave deposits. There are dubious reports in early newspapers. See Prehistoric Mammals. Felis concolor (= Puma concolor) probably the race coryi. Mountain lion or cougar. The native lion Felis concolor schorgeri was eradicated in the last century. Unauthorized releases of cougars, likely the Florida cougar, have occurred. Department of Natural Re- sources biologist Adrian Wydeven records all Felis concolor. Woodcut. Joseph B. Holder. The Mus. reports annually, so far dubious, invalid, and Nat. Hist., by Sir John Richardson et al. 1877. possibly valid. Jackson (1961) reviewed early

REINTRODUCED AND EXTIRPATED MAMMALS OF WISCONSIN 59 records. One record I investigated that may Rangifer tarandus sylvestris (Richardson, have led to several of the reports in the Rich- 1829). Caribou. There are no records of this land Center and Blue River areas probably species, which may have been present and erad- was a release of one or both of a pair brought icated about 1910 (Jackson, 1961). The spe- from Florida and fed on road-killed deer. The cies seems to have persisted in northern Min- owners left the state separately, and the cou- nesota. A possibly valid record for Wisconsin is gars did not leave with them. Observations given by Schoolcraft (1831). There are some were frequent in the region; these were among records of a larger bodied strain of caribou, from the first reports of cougars thought to have Prehistoric Wisconsin (see that Section). returned to Wisconsin. Bison bison bison. North American Bi- Cervus elaphus nelsoni. A reintroduced son or American Buffalo. Eradicated. Bison species, in spring, 1995, into the Clam Lake ranged into southwest and southern Wiscon- Area, were 25 individuals, some that died, sin on the prairies and savannas. Their status but some are producing calves under strict was reviewed by Jackson (1961), who report- protection. This race might replace in remote ed that the last of them was killed by “Sioux areas our extirpated Cervus elaphus ca- Indians” in 1832. In the display at Kettle nadensis. The Yellowstone-Jackson Hole Moraine State Forest there is a buffalo robe herd that provided so many reintroductions donated by James Brom (“from Lower He- of elk in America (see Long, 1965) provided bron Road”, ostensibly shot by John Mall’s at least one in Wisconsin (at Trout Lake in grandfather in the early 1840’s. It was shot 1913 and 1917). These elk were reportedly on Mall’s farm near Rome Pond [Jefferson in poor condition, but the photos in Jackson Co.]. If this record is valid (which seems like- (1961) are of healthy Montana stock. These ly) then the buffalo survived in some small introduced elk did not range out of Vilas and numbers nearly ten years longer than Jack- Oneida counties, and did not survive later than son thought, and this robe, preserved (“rolled 1950. The Jerome Hunting and Fishing Club up”) in Mall’s barn, is the only known bison at Mercer, in Iron County, near Trade Lake, specimen from Wisconsin from the pioneer used elk from Montana, and failed to estab- stock. Bison have been reintroduced into sev- lish elk. In 1995, the governor T. Thompson eral protected areas, such as the Sand Hills and others persuaded the Department of Nat- Demonstration Area near Babcock, and to ural Resources to make another attempt. numerous game farms. Today there is a grow- Cervus elaphus canadensis. American ing number of people undertaking to raise Elk, Wapiti. Jackson (1961) reviewed the his- bison as domestic beef animals. This is a great torical records, and there are numerous ant- example of what can happen to an animal lers from prehistoric Wisconsin (see that Sec- once nearly extinct, but through extensive tion). He believed this race of elk was eradi- efforts of William T. Hornaday and his sup- cated by 1875. It formerly was wide-spread. porters was saved from extinction (see Long, Alces alces. Moose. American Moose. 1965). Now it is useful to humankind again. The moose was probably eradicated at the Hybrids called “cattalo” may be obtained turn of the century, with the last record near by crossing Bison and domestic cattle. Some the Minnesota border in 1921 (see Jack- mammalogists refer Bison to the genus Bos. son, 1961). Some small groups may have I note the hybridization is not successful in survived in remote northern marshes. To- some crosses and the hybrid is obviously less day there are breeding records nearly every viable than the bison. Therefore, I am not year for moose that apparently reintroduced persuaded to consider the bison in Europe themselves in northern Wisconsin. See ac- and North America, with their many fossil count of that species. species, as conspecific with domestic cattle.

60 THE WILD MAMMALS OF WISCONSIN Felis catus. The house cat is not treated cats, which show rapid turnover, especially in this book as a wild species. It is more do- of males. Some diseases are transmitted to mestic than, say the house mouse (Mus mus- humankind by cats, and Toxoplasma is a real culus), although feral cats are important and threat to the offspring of pregnant humans. in many situations (especially in the southern Exotics. Some exotics and interesting counties) as self-sufficient as the mice. If hu- records that I have learned about in Wiscon- mans would cease partially feeding and re- sin include a prairie dog living at least 8 years newing the numbers of feral cats (by releases and keeping too many), probably the feral cats would soon disappear. House mice require no care from humans except in laboratories. John Coleman (1995) and Coleman and Temple (1993) found that farm-dwelling house cats (Felis catus), more or less feral and especially in southeast Wisconsin, make a huge impact in ecosystems by predation on songbirds, and even more on small wild mammals. The feral and domestic cats together may be the major predator in the state, especially in the agricultural and urban habitats. Coleman estimated that over 14,000,000 Wisconsin songbirds were killed annually by free-ranging cats, and double that number must be added to the predation by city and rural pets. The incredible total should make every cat loving naturalist pause for thought, 14 million song birds in only one state. Many more small mammals than that are killed, although these may be less beneficial. The average number of farm cats was between 8-9 cats, American elk (= Wapiti). Tho(?) Lundeen,1835, Cuvier’s and at non-farm rural homes about 4 to 5 Animal Kingdom, 1863. cats. The presence of up to 35 cats at a single farmstead was not uncommon. Farmers seemed overwhelmed by their own cats, hoping someone would help with their disposal. Farmers limit the numbers of cats by how much they feed them, but usually provide only sufficient food for approximately five. Ten cats survive on the rations for five. Each hungry cat may kill as many as 100-600 birds locally per year, as well as many small mammals. Although unwanted cats on farms may be shot, few (only 13 percent) are neutered. Compared to a density of about 34 raccoons, five skunks, and 18 opossums per square mile, the density of cats, at 104 per square mile, Sika deer Japanese exotic. Flower and Lydekker, 1891. seems incredible. Cars and diseases kill many By Lord Powerscourt.

REINTRODUCED AND EXTIRPATED MAMMALS OF WISCONSIN 61 in southwest Wisconsin (brought to my atten- crosorex) hoyi (holotype, see Long, 1972a). tion by Scott R. Craven), and one nutria near However, the species S. fumeus reportedly Stevens Point (1963) (see Long, 1970). Sika (but never verified) recently invaded northeast deer (Cervus nippon) escaped or were re- Minnesota, near the northwest Wisconsin leased from a game farm in southeast Wis- border, circling around the north shore of consin, and, reportedly, Fallow deer were re- Lake Superior. I have not seen the specimens leased as well, where hunters have reported (see Jannett and Oehlenschlager, 1994). The seeing adult deer with spots. Sika deer are species is clearly recognizable by flattened smaller than either Fallow or white-tailed deer, skull, dark venter, and size intermediate be- and have slender antlers in the males. In ad- tween the masked and arctic shrews, which dition to white spotting, there is a dark mane have high arched crania and mistakenly have in winter and a conspicuous white rump patch. been said to resemble fumeus closely. The The fallow Deer (Dama dama) is probably tail exceeds 38 mm in length. eradicated. It can be identified by larger body Perognathus flavescens Plains Pocket size and flattened palmate antlers (in males Mouse. This heteromyid, having external only) having numerous points. Wydeven and cheek pouches and elongate hind limbs, has Wiedenhoef (2002) record the odyssey of a four upper cheek teeth on each side, the dor- wolverine that escaped from a game farm in sum is cinnamon buff, the venter creamy Marquette County, apparently was observed white, and the upper incisors are grooved. It several times in the wild, and likely was the inhabits dry sandy prairies across the Missis- wolverine killed on the road in Rock County sippi River in Minnesota, but is rare there. in December 2001. Microtus chrotorrhinus Rock Vole. Known from one record in northeast Minne- sota. The vole is boreal and might have lin- HYPOTHETICAL LIST OF MAMMALS gered in Wisconsin in some boreal, rocky POSSIBLY OCCURRING IN OR INVADING habitat. It has an orange patch on the nose, WISCONSIN and tends to yellowish brown dorsally, whereas the venter is white. Some mammals that may in the future be Microtus xanthognathus. Yellow-cheeked found in Wisconsin may have been here all Vole. Recorded under the name Arvicola xan- the time, undetected, or they may invade the thognathus, yellow-cheeked vole, by Lapham state if habitats become favorable for range (1853: 340). The locality Racine would indi- dispersion. Most of the mammals listed here cate the record was based on Hoy’s notes. The are nearby or mobile enough to invade Wis- species is unknown from Wisconsin. consin. Phenacomys intermedius. The Heath- Sorex fumeus fumeus Miller, 1895. A er Vole may occur in northwest Wisconsin dubious record is the U. S. National Museum on sandy grassland or dunes. It occurs in specimen from southeastern Wisconsin, sent northeast Minnesota, and I have collected it from Racine by Dr. P. R. Hoy, in 1853. There in Ontario a short distance north of Lake are no other Wisconsin records, nor do the Superior. The cheek teeth are dark black- habitats at Racine seem appropriate. The ish, and especially in the lower jaw show deep known geographic range of the species was re-entrant angles that almost cut through the hundreds of miles away (Kentucky and north- teeth. The nose and rump are tan, and the ern Ohio). Probably the Wisconsin locality was venter is white. that of Hoy, not of his specimen. A mistaken Rattus rattus. Roof Rat or Climbing Rat. Racine locality was likely also for Microtus This long and scaly tailed rat, also called the xanthognathus (see below) and Sorex (Mi- black rat, regardless of its color, will likely be

62 THE WILD MAMMALS OF WISCONSIN found in Milwaukee, Green Bay, or at some VISITANT BATS port city on Lake Michigan, establishing itself briefly as an introduced rat. It was once taken Transported by winds, and often flying in long in Chicago. The tail is longer than half the dispersals of young-of-the-year, various bats total length, but see account of one long-tailed often are found far from their normal breed- rat under Rattus norvegicus. ing ranges. Those taken near Wisconsin in- Antilocapra americana. Pronghorn. By clude the Mexican free-tailed bat, Tadarida the name Antilope americana, included with brasiliensis, which has the tail extending well the Wisconsin mammals by Lapham (1852: beyond the uropatagium; the small-footed 44; 1853: 340), who cites only “N. W. Terri- Myotis, Myotis leibii, which is yellowish, has tory, (Mr. Say)”. Hoy (1882: 255) concurred, a dark mask on its face, and the calcar is believing the pronghorn ranged as far east as keeled; the gray bat Myotis grisescens, which Lake Michigan”. There is no evidence the has gray color all the way from the tips of the pronghorn ever occurred in Wisconsin, al- hair to the bases, and the wing membrane though ranging into Iowa. attaches to the ankle above the foot.

REINTRODUCED AND EXTIRPATED MAMMALS OF WISCONSIN 63 CHECK-LIST OF WISCONSIN’S Sylvilagus floridanus Eastern WILD MAMMALS Cottontail...... 150

With the names of orders and families Order Rodentia Sciuridae Order Marsupialia Marmota monax Woodchuck or Didelphidae Groundhog...... 158 Didelphis virginiana Virginia Opossum .... 69 Spermophilus tridecemlineatus Thirteen-lined Ground Squirrel ...... 164 Order Insectivora Spermophilus franklinii Franklin’s Talpidae Ground Squirrel ...... 171 Condylura cristata Star-nosed Mole ...... 76 Eutamias minimus Least Chipmunk ..... 175 Scalopus aquaticus Eastern or Tamias striatus Eastern Chipmunk ...... 180 Prairie Mole ...... 81 Tamiasciurus hudsonicus Red Squirrel ...... 187 Soricidae Sciurus carolinensis Gray Squirrel ...... 193 Sorex cinereus Masked or Cinereous Sciurus niger Fox Squirrel...... 203 Shrew ...... 86 Glaucomys sabrinus Northern Sorex arcticus Arctic Shrew ...... 89 Flying Squirrel ...... 210 Sorex palustris Water Shrew ...... 93 Glaucomys volans Southern Sorex (Microsorex) hoyi Pygmy Flying Squirrel ...... 214 Shrew ...... 97 Blarina brevicauda Northern Castoridae Short-tailed Shrew ...... 101 Castor canadensis American Beaver ... 220 Cryptotis parva (eradicated?) Least Shrew ...... 106 Geomyidae Geomys bursarius Plains Pocket Order Chiroptera Gopher ...... 229 Vespertilionidae Myotis keenii (= Myotis septentrionalis) Muridae Keen’s Myotis or Keen’s Bat ...... 114 Reithrodontomys megalotis Western Myotis lucifugus Little Brown Bat ...... 117 Harvest Mouse ...... 236 Myotis sodalis Indiana or Social Bat .... 121 Peromyscus leucopus Lasionycteris noctivagans White-footed Mouse ...... 241 Silver-haired Bat ...... 124 Peromyscus maniculatus (maniculatus Perimyotis subflavus Georgian and bairdii) Forest and Pipistrelle ...... 127 Prairie Deer Mice ...... 252, 257 Nycticeius humeralis Evening Bat ...... 130 Clethrionomys gapperi Red-backed Eptesicus fuscus Big Brown Bat ...... 131 Vole ...... 262 Lasiurus borealis Eastern Red Bat...... 135 Pitymys ochrogaster (= Microtus Lasiurus cinereus Hoary Bat...... 138 ochrogaster) Prairie Vole...... 267 Pitymys pinetorum (= Microtus Order Lagomorpha pinetorum) Pine Vole ...... 274 Leporidae Microtus pennsylvanicus Lepus townsendii (eradicated?) Meadow Vole ...... 280 White-tailed Jack Rabbit ...... 144 Synaptomys cooperi Southern Bog Lepus americanus Snowshoe Hare ..... 146 Lemming ...... 286

64 THE WILD MAMMALS OF WISCONSIN Ondatra zibethicus Muskrat ...... 292 Mephitis mephitis Striped Skunk ...... 404 Rattus norvegicus (introduced) Spilogale putorius (eradicated?) Norway Rat ...... 299 Eastern Spotted Skunk ...... 410 Mus musculus (introduced) House Lontra canadensis (= Lutra canadensis) Mouse ...... 301 River Otter ...... 414 Taxidea taxus North American Zapodidae Badger ...... 421 Zapus hudsonius Meadow Jumping Mouse ...... 305 Felidae Napaeozapus insignis Woodland Felis concolor (= Panthera concolor or Jumping Mouse ...... 311 Puma cougar) (eradicated) Mountain Lion or Cougar Erethizontidae Lynx canadensis (nearly eradicated) Erethizon dorsatum North American Canada Lynx...... 428 Porcupine ...... 316 Lynx rufus Bobcat ...... 433

Order Carnivora Order Artiodactyla Canidae Cervidae Canis latrans Coyote ...... 323 Cervus elaphus (eradicated, now partially Canis lupus (reintroduced, from Minne- domesticated) Elk or Wapiti sota?) Timber Wolf or Gray Wolf .... 330 Odocoileus virginianus White-tailed Vulpes vulpes Red Fox ...... 339 Deer ...... 440 Urocyon cinereoargenteus Gray Fox .... 345 Alces alces (eradicated?, re-colonized from Minnesota) Moose ...... 450 Ursidae Rangifer tarandus (eradicated) Ursus americanus Black Bear ...... 351 Caribou

Procyonidae Bovidae Procyon lotor Raccoon ...... 359 Bison bison (eradicated, now partially domesticated) Bison or American Buffalo Mustelidae Martes pennanti (reintroduced) Fisher ...... 369 A “Trivia” question: What Wisconsin mam- Martes americana (reintroduced) Pine mals are known by fewer than five reports or speci- Marten or American Marten ...... 375 mens today? Least Shrew, Indiana Bat, Evening Bat, Martes foina (introduced) Stone White-tailed Jack Rabbit, Franklin’s Ground Squirrel, Marten ...... 378 Stone Marten, Spotted Skunk, Canada Lynx, and sev- Mustela nivalis Least Weasel ...... 382 eral kinds eradicated long ago. Mustela erminea Ermine...... 387 Mustela frenata Long-tailed Weasel .... 392 Neovison vison Mink ...... 398 Gulo gulo (Prehistoric or eradicated) Wolverine*

* Gulo gulo. Jackson (1961) recognized the wolverine Gulo luscus (= Gulo gulo) as a Recent mammal; it is no longer in the state and perhaps eradicated. Possibly this mammal was prehistoric.

CHECK-LIST OF WISCONSIN’S WILD MAMMALS 65 KEY TO ORDERS AND FAMILIES 5B' Infraorbital canal narrower than foramen OF WISCONSIN MAMMALS magnum, quills lacking ...... 5C 5C Tail flattened dorsoventrally, hind feet 1 Incisor teeth 1-5/1-4 (= 5/4) on each webbed, cheek teeth each having 10-12 side; marsupium present in females, epi- transverse ridges ..... Beaver Castoridae pubic bones present, hallux opposable 5C' Tail not flattened dorsoventrally, hind feet to hind toes ...... Order Marsupialia: not webbed, teeth lacking 10-12 trans- Didelphidae Opossum verse ridges ...... 5D 1' Incisor teeth not 5/4, marsupium and 5D Upper cheek teeth number more than epipubic bones lacking, hallux never op- three, or in Napaeozapus an elongate posable to toes ...... 2 tail is tipped with white ...... 5E 2 Flight membrane comprised of finger 5D' Upper cheek teeth number three, tail not webbing, fingers elongated but claws lost tipped with white ...... from disuse excepting that of thumb ...... Rats and mice Muridae ...... Order Chiroptera: 5E Postorbital processes absent, tail long and Vespertilionidae Bats scaly, hind feet elongate ...... 2' Flight membrane lacking except lateral ...... Jumping mice Zapodidae membrane between wrist and pes present 5E' Postorbital processes present, tail fairly in Glaucomys, claws or hooves present long to elongate and always well-haired on digits ...... 3 or bushy ...... Squirrels Sciuridae 3 Upper incisors absent, digits provided 6 Fur on dorsum short and felt-like, zygo- with hooves ...... Order Artiodactyla: ma lacking or exceedingly slender, pre- Cervidae Deer maxillary-maxillary suture fused and in- 3' Upper incisors present, digits having distinct or if present manus is much larg- claws ...... 4 er than hind foot ...... Order Insectivora 4. Diastema present behind gnawing inci- 6A Manus larger than hind foot, zygoma sors ...... 5 slender ...... Moles Talpidae 4' Diastema lacking in tooth row ...... 6 6A' Manus smaller than pes, zygoma lacking 5 Incisors 1-2/ 1 (=2/1) but the second is a ...... Shrews Soricidae minute peg, hind limbs elongated for 6' Fur long on dorsum, especially guard saltatorial locomotion, pinnae of ears hairs, zygoma deep and functional as elongated ...... Order Lagomorpha: origin of masseter muscle, premaxillary- Leporidae Hares and rabbits maxillary suture distinct, and canine tooth 5' Incisors 1/1, hind limbs and ears usually prominent and anteriormost in the max- short ...... Order Rodentia: illary bone ...... Order Carnivora Sciuridae, Geomyidae, Castoridae, 6'A Four toes on hind foot ...... 6B Muridae, Zapodidae (this group 6B Only one upper (vestigial) molar present has long hind legs but short ears), on each side, retractible claws, five toes and Erethizontidae on front foot ...... Cats Felidae 5A External cheek pouches present, front 6B' More than one upper molar present, feet larger than hind feet ...... claws never retractible, four functional ...... Pocket gopher Geomyidae toes on front foot ...... Dogs Canidae 5A' External cheek pouches lacking, front 6'A' Five toes on each hind foot ...... 6C feet smaller than hind feet ...... 5B 6C Cheek teeth bunodont, carnassials not 5B Infraorbital canal wider than foramen adapted to shearing ...... 6D magnum, quills prominent on dorsum 6D Tail long, annulated, only two lower mo- and tail ...... Porcupine Erethizontidae lars, head of head and body less than 41

66 THE WILD MAMMALS OF WISCONSIN inches (= 1027 mm) ...... other is quantitative—the number of species in a spec- ...... Raccoon Procyonidae ified region, or species richness.” — Robert S. Hoff- 6D' Tail short, never annulated, three lower mann. . . . The Value of Abundance, Western Wild- molars present, length of head and body lands, 1991. exceeds 42 inches ...... Bear Ursidae 6C' Cheek teeth not bunodont, carnassials Wisconsin has been blessed with both a adapted for shearing ...... Mustelidae grand variety of mammals, from pygmy shrews to bears and moose, from the primitive opossum to the incredible star-nosed On mammalian biodiversity mole, and, of course, the courageous and fossorial badger. Especially along the “tension “There are several definitions of biodiversity. One is zone” between boreal and southern forms, qualitative—the variety of life forms, the ecological there is a fantastic richness of species coex- roles they perform, and the genetic diversity . . . An- isting together so long as humans permit.

Photo of opossum Didelphis virginiana. By John E. Long

KEY TO ORDERS AND FAMILIES OF WISCONSIN MAMMALS 67 TAXONOMIC ACCOUNTS OF include the “order” Polyprotodontia Kirsch, MAMMALS OF WISCONSIN or a quite different grouping, Marsupicarnivo- ARRANGED IN ORDERS, ra Ride. American Didelphids do not proper- FAMILIES, GENERA, SPECIES ly belong to any grouping of Australian mar- AND RACES supials, nor even with the South American Microbotheriidae (tiny Dromiciops australis) “One of the marvels of early Wisconsin was the from Chile (unless this little family is ancestral Round River, a river that flowed into itself, and thus or nearly so to American marsupials on the sped around and around in a never-ending circuit. one hand and to all Australian marsupials Paul Bunyan discovered it.... Wisconsin not only branching off the line separately). A possibil- had a Round River, Wisconsin is one. The current ity might be that Dromiciops is separate from is the stream of energy which flows out of the soil both American and Australian assemblages. [and sun] into plants, thence into animals, thence All the American opossums, Didelphidae, back into the soil.... Ecology is destined to be the seem allied by foot morphology, biochemical lore of Round River. Diversity [of plants and resemblances, and sperm-pairs (except little animals] means a food chain aimed to harmonize Dromiciops). Marshall et al. (1990) placed this the wild and the tame in stability, productivity, and little marsupial in a separate “order” Micro- beauty.” — Aldo Leopold, Round River, 1953. biotheria. This arrangement is not much different from the classical classification (of George Gaylord Simpson), of a marsupial Order MARSUPIALIA superfamily Didelphoidea containing Didel- phidae and Microbiotheriidae. The Marsupialia is one of the most ancient groups of extant mammals. In earlier ages it inhabited all the major continents, but today Family DIDELPHIDAE Gray it is confined to the Americas and the Austra- lian region. The marsupials are mammals The fossil record reveals that marsupials were known by their marsupium, a pouch in which generally replaced on one continent after an- the female carries and protects her young. other by the advent of modern placental mam- However, other mammals have pouches, and mals (which all have villi rooting the chorio- some marsupials lack them. No other mam- allantoic placenta into the flesh of the uterus). mals in North America have “epipubic bones”, The Didelphidae has become an exception to also called “marsupial bones”, one attached this geographic replacement, having success- to each pubic bone. Marsupials are grouped fully invaded (or reinvaded) North America af- because of common ancestry in an ancient lineage. They show much adaptive radiation, and taxonomists have split the group into subgroupings that are problematic and subjective. Many taxonomists today regard marsupials as a superorder, or even a cohort, comprised of several subgroupings formerly known as families considered to be orders themselves. One such is Didelphimorphia. Didelphidae, including the Wisconsin opossum, have been included with thylacines and other Australian marsupials. Such sub- marsupial categories (orders, or suborders) Opossum woodcut. [J. G.] Wood’s Nat. Hist. 1880’s.

68 THE WILD MAMMALS OF WISCONSIN ter the Pliocene Epoch. This American family, the state of Virginia. R. H. Baker (1983), quot- existing since the Cretaceous Time Period in ed the early settler John Smith’s note that North America, shows little evolutionary “opossum” is derived, from the Algonquian change for some 70,000,000 years of evolu- “apasum.” The name Didelphis marsupialis, tion. Didelphis virginiana is the only marsu- formerly applied to this species, is now restrict- pial species found in the temperate habitats of ed to the Central American opossum (Gard- the United States and southern Canada. It is ner, 1973), which differs significantly from our still expanding its range northward from its species in chromosomal pattern (fundamental South American origin (Gardner, 1982). number 20 instead of 32).

Genus Didelphis Linnaeus Didelphis virginiana virginiana Kerr Opossums 1792. Didelphis virginiana.The animal kingdom... “The brain ... indicated by the structure of the skull p. 193. Type locality Virginia. is remarkable for the proportionately large size of 1952. Didelphis marsupialis virginiana: Hall and the olfactory lobes, and the small size of the Kelson. Univ. Kansas Publ., Mus. Nat. cerebral hemispheres; these contracted in front and Hist.,5: 322. destitute of convolutions. As compared with the 1961. Didelphis marsupialis virginiana: Jackson, brain of the Mammalia ... the opossum furnishes Mammals of Wisconsin, p.17. the most remarkable contrast in its small size in proportion to the bulk of the animal. [However] ... Both of the aforementioned species the young return to the pouch to suckle and when names have been used by numerous Wiscon- danger threatens. During this time the female ... will sin workers. suffer any torture rather than permit the pouch to Description. The opossum is about the be opened.” — G. B. Waterhouse, 1848, Natural size of a house cat, muzzle slender, eyes con- History of the Mammalia, Vol. 1. spicuous and dark, fur shaggy and grayish, paler below where guard hairs are sparse, ears The description for the genus in Wiscon- rounded and paper thin, tail scaly (with clus- sin is written in the description for the sub- ters of ancient tactile hairs called “Dreiartgrup- species Didelphis virginiana virginiana Kerr pen” projecting outward from among the (see below). scales), large toe (hallux) on hind foot thumb- like, set apart from other toes (opposing them) and lacking a claw or nail. The plantar sur- Didelphis virginiana Kerr faces show friction ridges for climbing. Skull Virginia Opossum with prominent sagittal crest, small braincase, numerous incisors (5/4 = 18), prominent ca- The name Didelphis was proposed by Linnae- nines, and tribosphenic (trituberculate) molars. us. The word, literally “double love”, possibly The nasal bones widen posteriorly. There are means two penises, for the penis is forked. two pairs of palatine vacuities. No other Wis- There are also two uteri, which is what the consin species has marsupial (epipubic) bones Greek Dis + Delphus means according to on the pelvic girdle, nor a marsupium in the Waterhouse (1848). The name doubtfully re- female. There are five toes on each foot. Males fers to the double womb in time, i.e., an inter- possess a pendulous prepenile scrotum, and nal one for the embryos and an external pouch a urogenital sinus connects to the rectum in in which undeveloped young are nourished either sex. There are anal scent glands in the following birth. The name virginiana refers to rectum used for defense. The paired vaginae

TAXONOMIC ACCOUNTS / ORDER MARSUPIALIA 69 Skull of opossum.

blackish. A typical fur is mottled gray and whitish, the face whitish and feet blackish (with white toes). The eye-ring, eyes, and the inner surface of each ear flap are black. Yel- lowish stains on the chest (from skin glands) Opossum’s left hind foot. After Jackson. are seen in males. A winter-killed albino was found in Stevens Point. Hartman(1952) called this color phase “albinotic” because the eyes are black. Although Jackson (1961) mentioned that females are larger, Gardner (1973) found males are significantly larger, in 11 measurements. Hamilton (1958) found males about 32 percent heavier. Some standard measurements vary from a mean 24-35 in (609-900 mm) for total length, tail length 10-15 in (approximately 242-381 mm), hind foot 2-3 in (50-76 mm), and ear length about 2 in (40- Photo of mother and four young riding in the dorsal fur. F. M. Blake & J. Spero. Some deny that this happens. 50 mm). The greatest length of the skull varies from 80-127 mm, and weights from 4 to 15 pounds (1.8-6.8 kg). Weights reported combine as one that opens into the urogeni- from nearby states are given in the table Mar- tal sinus distally and at the other end, near 1. A 14-pound old male would be exception- the time of birth, to a temporary birth canal ally large. About a third and even more of the for the young. In males and females the uro- weight may be stored body fat, to endure the genital sinus and rectum although separate winter. Cleven (1988) gives the length as up are surrounded by a common sphincter mus- to 36 inches (92 cm), with weights to 12 cle. There are sweat glands on the feet and pounds (5.5 kg), and females up to seven tail. Chromosomes number 2N = 22, the Y pounds (3.2 kg). small, the fundamental number (i.e., contain- Dental formula. I 5/4, C 1/1, P 3/3, M ing elements of biarmed and acrocentric chro- 4/4 = 50. Young opossums may have mosomes) is 32 (Gardner, 1973). unerupted teeth. Fur in the opossum is grayish, but some The four molars above are primitive trig- specimens are white, dark brown, or even ons, the lower trigonids each has a talonid heel.

70 THE WILD MAMMALS OF WISCONSIN Geographic Range. The opossum is to be expected now throughout most of the state. It is not found on offshore islands in Lake Mich- igan and Lake Superior. On the Door Penin- sula it has been found only in recent years from north of the canal at Sturgeon Bay. See Map. Status. According to Jackson (1961) the opossum was known as early as the 1850’s in southern Wisconsin. It began rapidly expanding its range northward in the early 1920’s, possibly with the opening up of the North Woods (by the extensions of roadways, cutting of timber, etc.). Most records prior to 1961, and since then, are from southern

Maps showing geographic distribution of the Opossum in Wisconsin & North America.

TAXONOMIC ACCOUNTS / ORDER MARSUPIALIA 71 counties. Since 1965, specimens in the Uni- Lake Superior (Robinson and Heitman, versity of Wisconsin-Stevens Point museum, 1986). By 1996, opossums had advanced in a few from northern counties where no opos- western Wisconsin northward as far as Luck, sums were reported, permitted a crude, sin- Wisconsin. Another advance is to 9 mi. E gular estimate of the rateof dispersion (Long Fifield, in Price County (4 Nov. 1996). Re- and Copes, 1968). Other range extensions cently (2005) it extended the known range to for the Upper Peninsula have been reported northwestern Douglas County (see Records, (Baker, 1983), some that invite the question Ron Parmela). whether the estimated rate as calculated was The opossum is more than a curiosity of correct. However, in the sample used by Long nature, our only Wisconsin marsupial, because and Copes the dispersion was measured point its food habaits involve numerous food webs by point, representing new arrivals (accord- in our ecosystems. For one thing, it eats many ing to people that lived in those areas). The harmful insects. It also cleans up much carri- rates measured in two separate estimates were on. It has some fur value. On rare occasions a little more than four miles per year, in a it has killed domestic chickens. But today do- northward direction. Other records from the mestic chickens are seldom kept outside or eastern counties and in Upper Michigan may even in outside pens. be pioneers in a different advance. Along Lake In Wisconsin, the opossum is not legally Michigan, where frost is less severe, and opos- protected. However, hunting and trapping for sums are spared some of the frostbite of tails it and other furbearers are allowed only in and ears so common in central Wisconsin, late autumn or winter. Although the flesh is the opossum invaded Upper Michigan even edible, and savored in some areas of the south- faster than at the rate measured in central ern United States, and the fur is fairly luxuri- Wisconsin. Smith (1988) recently reported ous, not many are harvested and the pelt price three separate records a little northward in is low (compared to other furbearers). In Delta County, Michigan (1986-1987). Almost 1935-1943, the highest harvests of this spe- directly north of Menominee County, pene- cies in America exceeded 2,300,000 pelts, trating wilderness all the way to Lake Superi- but peaks dropped off to about 150,000 in or, the opossum was reported in 1984, on 1958-9 (Deems and Pursley 1978). Highway 41 at Marquette, only 1 km from The opossum builds a nest of leaves in a hollow snag, a crevice, in farm buildings, or even brushpiles. It may carry bundles of leaves to its Table Mar-1. Some weights (kg) of opossums from nearby states, based on data in Brocke (1970), Hamilton (1958), nest in its prehensile tail. It usually spends the Blumenthal and Kirkland (1976), Wiseman and Hendrick- winter in a burrow, often one abandoned by a son (1950), Pippitt (1976), and Lindsay (1960). Compiled by striped skunk. In its wanderings it may use many Gardner (1982). dens for day resting, often only a day. Location N & Sex Mean & Range Foods. Opossums are omnivorous, and feed on a wide variety of insects, other inver- Michigan 40 M 3.6 2.8-4.6 tebrates, mice, birds, worms, eggs, seeds, 12 F 2.4 2.0-3.2 New York 83 M 2.8 berries, soft fruits of plants, and even shoots 60 F 1.9 or leaves. They have varied fare of carrion Pennsylvania 10 M 3.4 2.2-4.0 (especially rabbits, and including other dead 18 F 2.4 1.8-3.1 road-killed opossums). Knudsen and Hale Iowa 5 M 3.1 (1970) examined the stomachs of 151 Wis- 10 F 1.8-3.1 Illinois 9 M 3.8 1.7-5.9 consin opossums from all months of the year, 7 F 2.4 1.8-3.1 and reported 65 different foods. See Table Indiana 11 M 5.4 Mar-2 for seasonal use. Mammals comprised

72 THE WILD MAMMALS OF WISCONSIN Table Mar-2. Seasonal use of important opossum foods unopened, crawling overhand like swimmers (values based on 5 percent of occurrences) studied in south- through the fur for a distance of approximately ern Wisconsin, 1953-1958. 3 inches (76 mm) (Hartman, 1920). Inside Food Winter Spring Summer Fall the pouch they each seize a nipple and hang on. Usually there are 13 nipples, as many as Earthworms — 31 54 35 Insects 14 11 29 31 17, in a horseshoe pattern. Not all the anteri- Mice 23 17 4 19 or nipples may produce milk, but approxi- Songbirds 6 19 25 15 mately 11, and sometimes 12 or 13 young Cottontail 29 9 — 19 become attached. The number of young that Frogs 3 10 25 12 survive is usually limited to only 7 or 8. For Garbage 16 9 4 — Snakes — 10 13 8 several days, if the young is detached from Apples 6 6 4 15 the nipple, it attempts again to seize a nip- Chicken 10 4 13 — ple. After three days the expansion of the dis- After Knudsen and Hale, 1970. tal nipple anchors the offspsring for some 50 to 60 days. The young weighs less than half a gram after several days. Thus, the mother’s 25% occurrence of the diet and 41% by nipple elongates during the period of lacta- weight. Most of the animals were decayed tion, and expands distally to help keep the carrion. Earthworms, insects, songbirds, frogs, young attached until they grow larger. When snakes and chickens were common summer detached from older offspring, the elongated foods. Other foods included cottontails, nipple facilitates nursing outside the pouch. shrews, moles, squirrels, muskrat, Norway rat, After 60 to 70 days, when the young dogs and cats, even swine, bobwhite, ducks, weigh about 25 g each, they leave the pouch pheasant, pigeon, screech owl, crow, cray- occasionally, returning for milk and shelter fish, snails, even fishes, apples, other fruits, until weaning. The young stay with the moth- grains, vegetables, mast, and garbage. The er as long as 3-4 months. Weaning is com- most amazing food is venomous pit viper pleted at approximately 90-100 days. The snakes, which cause no harm to opossums mother calls her offspring by a clicking noise, even when snake bites are successful and fangs similar to that of the male as he pursues a eject their poison (Gardner, 1982, Kilm- female. Often the opossum leaves her young on,1976, and Werner and Vick, 1977). in the den to forage for food. It is often said Reproduction. Opossums reportedly emphatically little opossums do not cling to mate lying side by side, by means of a bifid the mother’s fur, but see Fig., also Hamilton, penis inserted into the female’s vagina. Cop- 1939: fig. 59). Females may breed at six ulation is brief. Mating begins as early as Jan- months of age. Young males also might breed uary in northern states, but usually peaks lat- the first year. Most opossums breed first as er in March. Gestation is no longer than 13 yearlings. They seldom live beyond 2 years; days. The litter of partially developed young some apparently live four. may number as high as 22 or 25, but usually Opossums show a progressive irruption of 18 or less. A late litter was observed on 18 the dentition. Behind the canines the premo- July, 1993, in northern Door County. These lars and molars erupt roughly in this sequence: naked young were only 35-37 mm in crown- At 3 months, dp 2/2, M 0/0; at 4 months 3/ rump length. After birth, the reddish, naked, 3, 1/2; at 5-8.5 months, 3/3, 2/3; from 7-11 undeveloped young (length approximately 13 months, 3/3, 3/4; about 10 months or older, mm, weight 0.13 g) clamber quickly from the 3/3, 4/4 (the full adult complement) (see Gard- opening of the vagina anteriorly to the pouch, ner 1952). Mature opossums show tooth wear swinging their heads from side to side, eyes on the anterior molars before showing wear on

TAXONOMIC ACCOUNTS / ORDER MARSUPIALIA 73 all of them. Growth of the young is fairly regu- (51 ha). Younger or nonresident individuals lar (almost linear) the first year, by which time wander long distances, as far as 7 miles (11.2 sexual maturity is attained. km) in Missouri (Reynolds, 1945). Through one Mortality. Cars in Wisconsin kill innumer- winter month (January, 1996) I observed one able opossums. Dogs hunt them, hounds trail- old opossum (until it disappeared) that wan- ing them at night and often treeing them so dered nightly, within an area of no more than hunters may shoot them. Trappers take them 2 acres. Males have larger ranges than females, for their fur. Coyotes, foxes, badgers (proba- and juveniles smaller than either. bly), and great horned owls feed on them (Jack- Population density is known to be as high son 1961). The chief enemies are people, dogs, as one opossum per hectare in Illinois (Stieg- and automobiles, and long, cold winters (espe- litz and Klimstra 1962). Hunsaker (1977) found cially in underweight juveniles, less than 1400 density to be 0.26 / ha (0.2-1.16 / ha). g each, Gardner, 1982). Seidensticker et al. Opossums seldom live longer than 2-4 (1987) mention in this order, weather, nutri- years, but in captivity one lived six (Baker tion, and disease, with natural enemies owls, 1983). In a land so cold that the ears and tail hawks, coyotes, domestic dogs, raccoons, bob- freeze, the longevity is short (only 3-4 years cats and some large snakes (on young opos- according to Crandall, 1964), and the intelli- sums). Statistics given were cars (35%), para- gence of the animal questionable, then, when sites (17%), winter freezing (9 %), only 4 % all things are considered, it is surprising that due to trapping, and 35% unknown. opossums are expanding range. Jackson (1961) listed diseases and para- Remarks. Hair from the fur of opossums sites including tularemia, ticks (Dermacentor is used as nesting material by white-footed mice usually), fleas, and internal worms (Physa- and chickadees. Titmice were observed pull- loptera usually). Rabies virus can infect opos- ing hairs from a live opossum (Packard, 1949). sums, but usually there is little incidence. Sev- Small opossums hole up during freezing eral arboviruses infect opossums and could weather, but those larger than 4 kg changed pose a threat to other animals. Karl E. Shew- dens and moved about even then (Pippitt make, M.D., told this writer (1999) that opos- 1976). In winter the opossum sleeps a lot in sums present a problem as a carrier of a virus its more permanent winter den, but its body (called EPM) that causes a polio-like disease temperature does not appreciably drop. Year in horses in Wisconsin. It does not infect hu- round, it varies 32-38 degrees C, and ther- mans, and is transmitted by opossum feces. moregulation begins at 55 days (completed Horses eat the fecal material with meadow at 94 days). Opossums use their scaly, pre- grass or in hay, but are more likely to eat it hensile tail not only to clamber about in trees, mixed with horse food when the two mam- but to curl it around clumps of leaves to trans- mals share the food trough. Spotted fever, port them to their nests (dens). “Playing ‘pos- plague or tuberculosis organisms may infect sum,” of course, is feigning death, mouth opossums. Other diseases are listed in Addi- gaping, body limp and prostate, a physiolog- son et al. (1987) and Barr (1963). ically comatose state. I have seen it in a young Home range and Density. In Wisconsin, opossum, after it was attacked by our dog. Cleven (1988) mentioned home range of the Males make an open-mouthed threat in opossum as 20 to 40 acres (= 8-16 ha). Hun- encounters, screeching and weaving from side saker reported home ranges up to 20 ha. to side, fighting, and even killing the oppo- Gillette (1980) studied radio-tagged opossums nent on occasion. Females also gape and show in the warm seasons and found that the males agonistic threats to other adults, but aggres- wandered on average some 270 acres (108 sion ceases if the female is receptive to mat- ha). The home range for females was 127 acres ing. Such attitudes ensure a rather solitary life,

74 THE WILD MAMMALS OF WISCONSIN with opossums often avoiding one another. primitive ears (lacking much or all of the ento- Noises include clicking, growls, hissing, and tympanic and much of the ear pinna), the gen- in anger a loud screech. McManus (1974) italia and anus are in close proximity (almost reviews opossum behavior. resembling the cloaca of monotremes), and five Additional Natural History. Hartman toes are on each foot. The zygomata are ab- (1952), McManus (1974), and Gardner (1982) sent in shrews, and slender in moles. reviewed natural history for the opossum. The antemolar tooth-rows are divergent Geographic Variation. There is only one posteriorly, and the cheek teeth are typically race in Wisconsin. dilambodont (W-shaped). The anterior teeth Specimens examined. Total, 55. Adams, are procumbent, i.e., scalpal-like or forceps- Brown, Dane, Dodge, Door, Douglas*, like, on the one hand, or are tiny incisiform Green, Jefferson, Juneau, Kenosha, Ke- teeth and caniniform teeth, on the other. The waunee, Manitowoc, Milwaukee, Monroe, fur is short and felt-like in the shrews, and Oneida, Polk, Portage, Price, Racine, Rich- soft and fleecy in the moles. Although some land, Rock, Sheboygan, Trempealeau, Ver- shrews, and all moles, have white teeth, the non, Washington, Waukesha, Waupaca, shrews of Wisconsin show chestnut-brown Winnebago counties. tips, which more specifically tend to comprise Other Records. Jackson (1961) exam- an outer layer made of several sublayers, the ined 9 specimens from Wisconsin. Sauk Co.: outermost brownish. Each lower, anteriormost Leopold’s shack (Mossman, 1980, not tooth has almost a straight-edged brownish mapped). Michigan. Robinson and Heitman outer wall enclosing the inner, elongated white (1986) “On Hwy 41” at Marquette, 1 km from enamel crown, with humps of red-brown in Lake Superior. 1984. Roadkills between Mari- tandem in some species analogous to cusps, nette and Escanabe. and in Sorex arcticus showing anteriorly even an inner brown tipping on one hump. In these teeth, Blarina and Cryptotis have nearly Order INSECTIVORA straight-sided red-brownish walls. They show, The Insectivores as does Microsorex, accessory cuspules on the upper “unicusps”. The terms unicuspid, “In the dark and cold embrace of Mother Earth, away unicusp, even unicuspate are not technically from the cheering sunlight and the beautiful upper correct for such shrews. world that we enjoy, there dwells a group of mammals The Wisconsin insectivores belong to a so strange and yet so useful to man, that they excite distinctive suborder Lipotyphla. The other sub- our admiration.... Pass not unthinkingly the moles and order Menotyphla includes the African jump- shrews, for they have been most cunningly designed ing shrews and the Tupaiidae (except some to serve a definite and important function in the workers, including myself, assign Tupaiidae or economy of nature.”— W. T. Hornaday, in The tree shrews to the order Primates). In shrews American Natural History, 1904. W-shaped ectolophs are conspicuous. In moles, not far removed from the ancestral insectivo- The order Insectivora, as old as Creta- ran Leptictid pattern, the ectolophs are said to ceous Period, is found on all continents ex- be W-shaped. Moles seem primitive in molar cept Australia. These some 400 species are pattern, retain a slender zygoma, are about the comprised of medium-sized to tiny mammals same as shrews in construction of their audito- having elongate and pointed snouts, small eyes, ry bulla (some have the bulla complete by addition of an entotypanic expansion), and the den- * Hwy 13, Amnicon R., sect.10, May 9, 2006. Road- tition of Condylura has the primitive number kill reported by R. Perala. of teeth (44). Therefore, moles are discussed

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 75 before shrews, which is not conventional, but 1’ Snout devoid of tentacles, anteriomost in line with evolution of tiny size in shrews. upper teeth elongate, chisel-like and robust, tail short and not tapered...... Eastern or Prairie mole Key to Families of Scalopus aquaticus Wisconsin Insectivores

1 Size larger (total length exceeds 150 mm, Genus Condylura Illiger weight more than 30 g), external ear pin- Star-nosed Mole nae lacking, forefeet broad, twice the width of hind feet, zygomata slender but Condylura cristata cristata (Linnaeus) always present, teeth exceeding 2 in number ...... Talpidae, Moles 1758. Sorex cristatus Linnaeus. Systema Naturae, 1’ Size smaller (total length less than 150 ed. 10. Vol. 1, p. 53. mm, weight less than 30g), external ear 1819. Condylura cristata: Desmarest. J. de Phys., pinnae inconspicuous but present, fore- Chim., Hist. Nat., et des Arts. 89:230. feet small, hardly wider than hind feet, zygomata lacking, teeth 32 in number The name Condylura Illiger, 1811, is or fewer ...... Soricidae, Shrews Greek, and means knobbed tail based on a faulty drawing by De la Faille. Linnaeus provided the name cristata, which means little Family TALPIDAE Fischer crest, and is often featured in moles. External Moles measurements are given in Table Ins-1. Dental Formula. DF = I 3/3, C 1/1, P “These beasts are all blind... and therefore came the 4/4, M3/3 = 44, the so-called primitive num- proverb, Talpa caecior tuphlotaeros alpalacos, ber of placental mammals. Howard Whidden blinder than a Mole; to signifie a man without (personal communication) informs me the judgment... yet if any man look where the eies genus is known in the Pliocene of Poland, should grow he shall perceive a little passage, [and] supporting my view that the mole is primi- by drawing up the little skinne... therefore of them. — tive. Some premolars are triconodont (3 cusps Topsell, Historie of foure footed beastes, aligned) and do not occlude with their ana- London, 1607. logs in the opposite jaw. The canines are caniniform, which is not always the case in Chief characters are summarized in the moles. The I 2/ is so minute it may be ob- Key above. These fossorial mammals feed scure (see skull fig.).attributed to a peculiar mostly on earthworms (Oligochaeta) and bee- sagittal ridge, but much more likely refers to tle nymphs. the rosy-pink cluster of tentacles on the snout. In Latin, crista also means a tuft on the head. This corona of 22 bilaterally arranged tenta- Key to Wisconsin Moles cles, tactile in function, provides a basis for the name “star-nosed mole.” 1 Twenty-two tentacles surround nosepad Description. This mole has the typically (rhinarium), anteriormost upper teeth broad hands with robust claws characteristic spoon-like, elongate tail tapered at ante- of moles. They are narrower than in the East- rior and posterior ends ...... ern mole Scalopus. The star-nosed mole is ...... Star-nosed mole readily identified by the 22 pinkish or gray (in Condylura cristata dried skins) tentacles surrounding the two

76 THE WILD MAMMALS OF WISCONSIN Skull of the star-nosed mole.

Geographic Range. The star-nosed mole dwells in northern Wisconsin and Upper Mich- igan, restricted from most islands in Lake Mich- igan, and apparently from the Door Peninsula. Status. Condylura is uncommon, restricted to wet, boreal soils of swamps and stream Musculature and tendon structure of the nasal rays of valleys in northern counties. The status seems Condylura. After Grand et al. 1998. unchanged from the turn of the century, although the loss of wetlands and application nostrils, and a long, sparsely haired, fleshy of pesticides probably had an adverse effect. tail (tapered at either end). The eyes are small, The soils where this mole dwells are usually auricular openings prominent, feet have too wet for agriculture or building, and this scales, and 5 toes are on each foot. The fur is animal is seldom a nuisance. relatively coarse for a mole, dense and long, Habitats. This semi-aquatic mole is found with a softer underfur. Molts occur in June in black, wet soils, mucky humus, or sandy and October, when winter fur is attained. The loam, of stream and river valleys, swamps, skull is long and narrow, zygomata short and lake shores, and marshes mostly of northern slender, jaws narrow with widely-spaced (saw- counties. Occasionally it lives in lawns near like) antemolar teeth, molars mostly W-shaped water, in cedars, spruce, tamaracks, birches, with interior portion a shelf, premolars some- alders, jewelweed, marsh grasses, and sedg- what tricuspate, lower molars similar to up- es, often in association with jumping mice and per molars, but portion a shelf, premolars shrews. I have taken this mole (2) and Blari- somewhat tricuspate, M-shaped, auditory bul- na (2) from the same partially flooded tunnel lae incomplete. Chromosomes number 34, (Jordan Marsh, Portage County) on alterna- with a fundamental number of 64 (Petersen tive nights. Don Follen, Sr., trapped two Bla- and Yates, 1980). The karyotype is much dif- rina and two Condylura sharing the same ferent than in other moles. tunnel (Oct. 16-28, 1967) near Arpin, Wood The star-nosed mole is nearly black dor- County. Condylura swims under ice and sally, the venter a paler blackish brown, tail, moves about on snow (Jackson 1961; also feet and head concolor with body. The wrists see photo). are tan. Worn pelages are browner or gray- Star-nosed moles tunnel less extensively er. The sexes are similar in size, unusual in than the prairie mole Scalopus, and the ridg- moles. External measurements are given in es are usually less than two inches wide. Table Ins-1. Where surplus dirt is mounded up, in wet globs

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 77 and layers, the mole hill may extend above the vegetation to a height of six inches, and may be as much as two feet wide. In a series of mole hills found in farmyards in eastern Wood County, the huge hills resembled those of Scalopus. Small trails, almost trenched runways, extended on the soil surface to a small intermittent ditch of water. The nest is a loose wad of leaves, grass, or both, constructed in an ovate chamber nearly seven inches in diameter and three to five inches in height. The nest has been observed sited between tree roots and under a decayed stump. Audubon (Audubon and Bachman, 1842-

Maps showing geographic distribution of Condylura cristata in Wisconsin and North America.

78 THE WILD MAMMALS OF WISCONSIN 1846) found one made of withered grasses in tricity over non-electrical areas. Whether the an excavation under a stump to which a tun- Eimer’s organs, small tactile sensory organs nel was excavated. The nest is usually three on the tentacles, are responsible for this sixth to ten inches underground. As many as three sense is not clearly stated, but Gould et al. tunnels enter the nest; each is about 40-50 suggest there is a great density of these or- mm in diameter. Young have been found in gan -receptors around the center of each set the nest (Jackson, 1961; Schmidt, 1931; Au- (of eleven) rays. They suggest the disparate dubon, loc. cit.; Rust, 1966). size of the rays enhances sensory or electri- Foods. Star-nosed moles feed mostly on cal perception. terrestrial and aquatic worms, insects, other Catanla et al. (1993) identified “stripes” invertebrates, and to a lesser extent aquatic on each side of the neopallium of Condylu- vertebrates (frogs, fishes) (see Jackson, 1961). ra, which areas apparently have evolved for Don Follen found earthworms in a Wood the special afferent impulses from each set of County mole’s stomach (October, 1967). One eleven rays. Taken altogether the evidence kept in captivity was fed earthworms in a bowl supports an explanation of an amazing elec- of water daily. The mole dived in swimming tromechanical sense in Condylura used to about beneath the surface with its tentacles detect prey and to explore surfaces, such as actively searching. When the worm was en- the water surface. There are some students countered the male seized it, carried it from of moles who are unconvinced. No one doubts the water to a corner of the aquarium, and that this mole has some sort of incredible noisily ate it. A plastic bass bait closely re- sense (taste?) for detecting worms. sembling a worm was never seized by the Reproduction. Jackson (1961) suggests swimming mole unless the plastic worm was the adult star-nosed moles that pair in autumn rubbed with a live worm, whereupon the mole may mate, but probably not until winter. Males seized the bait and carried it out of the water. apparently breed in February to mid-March. One could hear gnashing of tiny teeth, sug- Gestation is believed about 45 days. Breed- gesting that somehow the mole had tasted ing reportedly extends from mid-March the juices of the live worm. Whether the ten- through April. There are usually six young, tacles function in the tasting is unknown, for but the number may vary from 3-7. Eadie and no one has ever searched for taste buds in the tentacles. Possibly the tongue is used in tasting the surface of a prey animal that was detected with the tentacles. Gould et al. (1993) described a new function of the oral tentacles, as electroreceptors.. This new sense was unknown in mammals except for its recent discovery in the Austra- lian platypus (Ornithorhynchos anatinus). Gould et al. found attacks of moles correlated with specific parts of the worms, especially the clitellum, where electroactivity from secretions was measured as particularly strong. Other electrical “hot spots” were likewise selected as targets by the moles. Data from electrified fields in tunnel-mazes suggested electroreception in the oral rays. Moles even selected simulated worm fields of elec- Condylura cristata from Portage County.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 79 Hamilton (1956) found a mean litter size of be found in ideal habitat. These concentra- 5.4. There are eight mammae. Naked and tions are likely family groups, and some have blind at birth, the young are pinkish and only doubted that the density is so great. In cen- the vibrissae are visible hairs. Partially devel- tral Wisconsin I have taken several together, oped tentacles are obvious even in develop- suggesting about 3-4 per acre, and nearby ing embryos. Newborn young are remarkably none at all were trapped. In their wanderings large, reportedly measuring 70-75 mm includ- by use of water (they even swim under ice) ing the tail (25 mm). The young are born in and tunnels in the snow they may disperse late April to mid-June (which may account from one habitat to another, and often find for the paucity of breeding records, because themselves in some farmer’s yard by access little field work is carried out in the North of an ephemeral ditch of water. Woods until winter’s end). There is but one Remarks. Large star-nosed moles (60 g) litter per year. In six to eight days short fur consume oxygen at a rate of 4.2-4.5 cc/g/ appears dorsally, and in two days more the hr. Juveniles had lower metabolism, about half abdomen is haired. When the young leave the as much (see Petersen and Yates, 1980). nest at about one month or more of age their Additional Natural History. Peterson sleek fur covers the body. They each weigh and Yates (1980) reviewed biology for about 30 grams. Mrs. John Pollock of Rhine- Condylura. lander found five young in a nest on May 3, Specimens examined. Total, 61. Ashland 1932 (Jackson 1961). Alan Long caught one (Outer Island, Apostle Islands, 1), Bayfield, by hand weighing only 20 g, in Whiting, Wis- Brown, Calumet (Kiel, 1), Clark, Dodge, Ke- consin, in early July. This mole may have been waunee, Jackson, Juneau, Manitowoc born in early June. Although small it was well (Branch River Country Club, 1, Point Beach furred (see Table Ins-1). State Forest, 1 UW), Marathon, Marinette, The teeth erupt before the young moles leave the nest. Jackson (1961) mentions a small sagittal crest appearing in old adults. Table Ins-1. External measurements and weights of Small moles in this collection yield measure- Condylura from Wisconsin. Specimens selected having weight ments given in Table Ins-1. data on the labels. Mortality. Carnivores that prey on star- No. Locality / Measurements Sex/ Weights g. nosed moles, usually catching them above ground, are dogs, red foxes, fishers, striped Adults 5971 Drummond M skunks, and house cats (Hamilton 1936, Mu- 184-75-28 46 rie 1936, Schmidt 1931, Timm 1975, Toner 3795 Westboro M 1956). Aquatic predators include large-mouth 168-70-23 37.1 bass, bullfrog, and mink (Christian 1977, Pine 1005 Eagle River M 1975). Mike Eber told me a northern pike ate 179-70-24 46.9 1753 Abbotsford M a star-nosed mole in Monroe County. Hawks 188-76-25 49 and owls take them (see Baker 1983) and even 63 Portage Co. F the great gray owl (3 specimens in this collec- 190-72-21 42.7 tion are from owl pellets from Ashland Coun- 3716 Wood Co. M ty). Ticks, mites, fleas, tapeworms, and round- 176-70-25.5 45.8 worms parasitize this mole (Hamilton 1931, Of Weaning Size 3107 Marinette Co. F Jameson 1949, Scharf and Stewart 1980, 77-70-26 30.1g (Oct.6, 1971) Timm 1975, Yates et al. 1979). 3895 Whiting ? imm. Home Range and Density. Jackson 141-53-25 Estimated wt < (1961) thought as many as 30/ acre might 20 g (July 11, 1974)

80 THE WILD MAMMALS OF WISCONSIN Jackson (1961) and others refer to it as the prairie mole, which is an excellent and meaningful name. Description. The eastern mole is rat- sized, stout, with huge hands and enormous foreclaws (used for digging and pushing soil), head short and almost conical, lacking visible ear pinna or conch, and the auricular openings are small. The eyes are tiny, present but Head and forefoot of Scalopus. Scratch sketch H. E. suspended in skin, covered with fused eye- Anthony. lids; the snout naked anteriorly with nostrils opening upward; tail short (about one fifth of total length), scantily haired so as to appear Oneida, Portage, Price, Rusk, Sheboygan, naked, feet all with five toes, all webbed, pel- Taylor, Vilas, Waupaca, Waushara, and age plush (velvety) dense and smooth. The Wood counties. skull has stout, large but slender zygomata, Michigan. Other record. Big Summer Is- complete auditory bullae, truncated nasals, land, Lake Michigan. Schoolcraft Co.: Seney and anteriormost teeth (probably I 1/) large 1 UMinn. and directed vertically (almost chisel-like as in rodents). Chromosomes 2N = 34, fundamental number reportedly 64 (Hoffmeister, 1989). Scalopus E. Geoffroy Saint-Hilaire The eastern mole is grayish black or brownish black, often washed with a silvery See Description for Scalopus aquaticus. sheen, usually browner in winter, occasionally with an orange or rusty sheen on venter (environmentally caused?). Feet are dirty whit- Scalopus aquaticus (Linnaeus) ish above, with naked gray skin below. No. Eastern or Prairie Mole UW 2040 taken June 1, 1969, in Crawford County, is an albino. Scalopus aquaticus machrinus (Rafinesque) Male eastern moles are slightly but significantly larger than females in most exter- 1832. Talpa machrina Rafinesque. Atlantic Joumal, nal and cranial measurements. Weights of 1:6 1. Type from Lexington, Kentucky. males are given in Table Ins-2. 1832. Talpa servicea Rafinesque. Atlantic Journal, 1:62. Type from Nicholasville or, Harrodsburg, Ky. 1842. Scalops argentatus Audubon and Bachman. J. Acad. Nat. Sci. Philadelphia, 8:292. Type, from southem Michigan. 1905. Scalopus aquaticus machrinus: Elliott. Field Columb. Mus. Publ., 105, Zool. Series, 6:470.

The name Scalopus comes from the Greek Skalops, to dig. The word “aquaticus” means, of course, aquatic, but this is a mis- nomer because the mole is not aquatic. This is not the only mole found in eastern United States, but it is common and wide spread. Skull of Scalopus aquaticus.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 81 Dental Formula. Reported often as I 3/ more flattened heaps of soil. Often found and 3, C 1/0, P 3/3, M 3/3 = 36. However, confused with pocket gophers, the tunnel ends there is no way to account for the tiny spi- are not plugged (by pushing soil upward and cules variably present in the lower antemolar out), nor are the hills of dirt thrown in some series, nor to know with certainty the serial particular direction, but raised upward and homology with the upper ones. Some ante- spread more in a circular pattern. Such tun- molars are spicules either reduced in size in nels in the topsoil seem like wrinkles in a lawn, their evolution, or perhaps were retained milk but in winters the tunnels lie deeper where teeth. Consequently, one often observes teeth the worms and beetle larvae are more likely in excess of 36, and their presumed homolo- to be found. This deeper series of tunnels, gy with, for example, Condylura, cannot be reportedly 8 to 24 inches (Jackson 1961) and confirmed. For practical purposes, the upper doubtlessly a great deal deeper in winters of series of teeth of the eastern mole can provi- deep frost, is the home of the eastern mole. sionally be regarded as 3, 1, 3, 3. Jackson reports that this mole digs in lawns Geographic Range. This prairie mole is 18 feet per hour, and in a night can create a confined to the prairie counties of southern ridge 50 or more yards in length. The nest is and western Wisconsin, ranging to the south- about 5-6 inches (127-152 mm) in diameter, ern marshes of Portage and Wood counties, usually at a depth of 13 to 18 inches. Some- and northwest to Burnett County along the times the nest is only 5-6 inches underground. west border. See Map. The nest is comprised of grass and rootlets, Status. Unchanged since the 19th cen- occasionally of leaves, and at times is practi- tury. This eastern (or prairie) mole is often a cally void of nesting materials. The nest is sit- nuisance in lawns, cemeteries, and golf cours- ed in an ovate chamber about 8 inches long es. It is beneficial in soil formation and in eat- with diameter about 5 inches. ing beetle nymphs (grubs). It eats earthworms Foods. Jackson (1961) and Hisaw (1923) (of no appreciable or practical loss) and corn considered the primary foods (80 percent) of seedlings on occasion. the eastern mole to be earthworms, grubs and Habitats. The eastern mole occurs basi- adult insects. The remainder is plant materi- cally in sandy loams, grasslands, prairies, road- al. On occasion, snails, centipedes, and milli- sides, pastures, railroad rights-of-way, lawns, pedes are eaten. Plants include rootlets and golf courses, cemeteries, and cultivated fields. seeds, and captive moles feed on a variety of Hoffmeister (1989) mentioned their occur- fruits and vegetables (summary in Baker rence in almost any friable soil in which the 1983). West (1914) and Whitaker and moles can push or dig dirt in their tunnels. At Schmeltz (1974) found comparable percent- the edge of their range in central Wisconsin, ages. The season and habitat are important the moles seem limited in their northward dis- according to West who found moles feeding persion by the extensive marshlands, which, on carpenter ants, and in May and June on after draining by ditches and allowed to de- emerging May beetles. One mole caught in velop as prairie (for prairie chicken manage- winter had eaten 150 ants. ment in some cases) become suitable for Reproduction. Conaway (1959) obtained moles. In western Wisconsin the moles are information from Dane County moles. They found both in the valleys and on the uplands, breed but once a year, and young-of-the-year often sandy where soil is established on the do not breed. Breeding commences as early summits (e.g., at Ellsworth, Wisconsin). as February or March and continues into April. The ridges or mole runs of the eastern Gestation is thought to be about 45 days, but mole usually are wider than those of the star- may be only one month (Jackson, 1961; Con- nosed mole, and the hills larger, wider and away, 1959). No parous females were found

82 THE WILD MAMMALS OF WISCONSIN in late March when breeding was at peak. Con- away notes a few pregnancies after 5 May; one was as late as 4 July. The naked young are large at birth (about 50 mm in length). The litter size varies from 1 to 5, usually 4. There is some resorption of embryos. Testes develop rapidly in December, but mating occurs several months later. The young are haired out in about 7-10 days. In about five weeks, the young are more than half the size of their mother (Yates and Schmidly 1978; Jackson 1961, Hoffmeister 1989). Mortality. Eastern moles have few enemies because they so seldom come out above

Maps showing geographic distribution of Scalopus aquaticus in Wisconsin & North America.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 83 ground. Hawks and owls eat them on occa- Other Records. Milwaukee Public Muse- sion, and foxes, coyotes, and domestic dogs um records. Pierce, Grant, Columbia, Sauk, kill them (Arlton 1936, Davis 1951, Wallace Adams, and Buffalo counties. Ellsworth, No 1950, Wilson 1938, Errington 1935, Murie specimen (mole hills observed). Nottingham 1936, Schofield 1960, Van Hyning 1931). Ridge, No specimens. 2 mi. SW Osseo, also Often the mole is rejected as food because of Osseo (No specimens, mole hills observed). its rank odor. Parasites include the louse Eu- Town of Grant, Prairie Chicken Area on Bue- haematopinus abormis, fleas (Ctenophthal- na Vista Marsh, T21 N, R7E (no specimen, mus and Corypsylla), mites, and intestinal hills observed). Clark (Schmidt, 1931). worms (Monoliformis, Filaria, and Spiroptera) (see Jackson 1961, Baker 1983). Home range and Density. Arlton (1936) Family SORICIDAE Linnaeus and Harvey (1976) found home ranges for Shrews the eastern mole from 0.86 acres to two acres (0.75 -8.0 ha) and 10.6 ha (for males) Shrews are small to tiny mammals (including or 2.7 ha (for females). Arlton found a some of the smallest mammals in the world) branching tunnel system covering an area with conical, elongate and flexible muzzles (or about 0.35 ha. Moles wander much more snouts), tiny eyes hardly visible, ear pinnae extensively than pocket gophers (Yates and small but usually evident, front feet smaller Schmidly, 1978). Baker (1983) suggests that than hind feet, tail well-haired with short hairs, 1-2 moles/acre (2-5/ha) are probably nor- each foot having five tiny toes, skull bird-like mal density. He disputes Jackson’s (1961) with ovate braincase and slender rostrum, and estimate of 8-10 per acre. zygomatic arch incomplete. Anteriorly it per- Additional Natural History. Yates and sists as a vestigium, the maxillary process, and Schmidly (1978) reviewed the biology of Sca- the jugal bone is lacking. The anteriormost lopus aquaticus. upper tooth is bicuspate, its lower analog in Geographic Variation. There is a cline the dentary remarkably elongated, and usual- (i.e., a gradual change over some wide area) in ly along each border slightly tetracuspidate. increasing size northward from Tennessee, but The functional points (apices) of the cusps and it is not continued into central Wisconsin. That crests are usually red-brown unless worn away, is because the eastern moles in southern Wis- the auditory bullae are undeveloped, the tym- consin are likely recent invaders from the south- panic bone ring-like, and the pelage felt-like east. Specimens examined. Total, 29. Adams, (short and dense). Burnett, Chippewa, Crawford, Dane, Eau Claire, Grant, Iowa, Juneau, Monroe, Por- tage, Richland, Sauk, Vernon, Waushara, Wood counties.

Table Ins-2. Body weights of male Scalopus from Wis- consin , after Conaway 1959.

Months N Mean sd Adult March-April 34 121.5 g 11.35 Adult May-July 27 120.6 14.10 Adult August-November6 117.6 5.15 Juvenile May-July 28 95.7 8.75 Sorex cinereus and the larger S. palustris. Spencer Juvenile August-November10 107.8 10.79 Fullerton Baird, 1858.

84 THE WILD MAMMALS OF WISCONSIN Shrews have high metabolism per gram Key to Wisconsin Species Of Shrews of body weight. In Blarina the metabolic rate is 3.18 cm2/g/hr, with heart rate of Note: Sorex fumeus ought to be confirmed, 740-760/minute, and a body temperature an unlikely member of the Wisconsin fauna, of 38o C. (Doremus 1965, Neal and Lust- and it is not treated herein. The locality of Ra- ick 1973). In smaller shrews the metabolic cine may as well refer to the home of the col- rate per gram of body weight is even high- lector Hoy as to the shrew. It resembles Sorex er, highest of all in the pygmy shrews Mi- cinereus, but is recognized by dark underparts, crosorex (Buckner 1964:260). Animals that flattened braincase truncated posteriorly, and have such high metabolism need to eat reg- inner lophs of unicuspate teeth less pigment- ularly. In winter they must eat to compen- ed. The skull is longer than 17.5 mm. sate for loss of body heat because their surface area to mass ratio is high. The dense 1 Tail short relative to the body length, less fur is efficient insulation, but a great deal of than 30 mm (1 1/4 in), more than a third protection is afforded the shrews by deep of the total length, external ear conch snow in the north (under which the temper- hidden in the fur of the head ...... 2 ature is warmer). 1’ Tail long relative to the body length, more Shrews are found in Wisconsin in soils than 30 mm and more than a third of having a high content of moisture (Sasse the total length,ear pinna tiny but visible 1978; Getz 1961d). Sorex arcticus prefers ...... 3 marshes and shrub communities, Blarina 2 Size larger (head and body more than brevicauda is at home in grasslands or 90 mm), tail 18-30 mm, mature skull ex- woodlands, Sorex palustris is restricted to ceeds 23.0 in length, 5 unicuspate teeth stream banks and wetlands, and Sorex (Mi- ...... Northern Short-tailed Shrew crosorex) hoyi is an enigma and, although Blarina brevicauda rare, found in diverse environments. Cryp- 2’ Size small, about 90 mm or less in head totis parva is probably eradicated from and body length, skull much less than 20 Wisconsin sandy prairies. mm, 4 unicuspate teeth .... Least shrew How to study shrews. The advent of pit- Cryptotis parva fall trapping, i.e., sinking empty 1/2 or 1 3 Third unicuspate tooth smaller than gallon cans into soil (the rim flush with the fourth ...... 4 soil surface, soil packed closely against it) in a 3’ Third unicuspate tooth larger than fourth variety of natural habitats (punching holes in ...... 5 the bottoms in wet soils so the cans will not 4 Size larger, total length 131 or more, pel- float up) has led to greatly increased catches age dark blackish or slate-gray dorsally, of the rarest and tiniest shrews. It is possible hind feet fringed with hairs, third unicus- to obtain density data, distributional data, and pate tooth somewhat smaller than fourth of course, more information on embryo ...... Water shrew counts, parasites, and so on. By checking the Sorex palustris cans regularly in the night, and leaving food 4’ Size small, total length approximately 91 in the cans (insects, worms, canned dog food), mm or less, hind foot tiny, never fringed, it is possible to capture shrews alive. That third unicuspate tooth a minute and in- could lead to capture-recapture information conspicuous disk squeezed between ad- if marked shrews are released. A wonderful joining teeth, anterior teeth (bicusps) opportunity exists to keep the rare shrews spaced apart with expanded medial tines alive in large enclosures with natural soils and ...... Pygmy Shrew native vegetation. Sorex (Microsorex) hoyi

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 85 5 Size small, total length 102 mm or less, length of skull less than 16.5 mm, unicuspate teeth even except diminutive peg-like fifth, rostrum exceptionally slender, cranium slightly arched ...... Masked Shrew or Cinereous Shrew Sorex cinereus 5’ Size medium, total length greater than 102 (106-121) mm in adults, length of skull exceeds 16.5 mm, unicuspate teeth Skull of Sorex cinereus. large, rostrum wide, cranium prominently arched, tricolored pelage (black dorsally, brown, pale brown) in winter pelage .... is not entirely appropriate, for the shrew is ...... Arctic Shrew brownish gray, but it is far more appropriate Sorex arcticus than “masked” shrew because there is no mask. Besides, Jackson’s classic work on shrews may allow him to set precedence on names for Genus Sorex Linnaeus shrews, based on his extensive work on them. Long-tailed Shrews Furthermore, the animals called masked shrews recently have been divided into several species. There are five unicuspate antemolar teeth and Description. This tiny mammal is one of the tail is relatively long. Mammae: 6. the smallest mammals in Wisconsin, perhaps a little larger than Wisconsin shrews of the species Sorex (Microsorex) hoyi. The masked Sorex cinereus Kerr shrew has a slender snout, and the rostrum of Masked or Cinereous Shrew the skull is correspondingly slender. The tail is relatively long. The bicuspate teeth are hardly 1792. Sorex arcticus cinereus Kerr. Animal Kingdom, separated by expanded tines, therefore nearly p. 206. Type from Fort Severn, Ontario, Canada adjoined in front, and the four anterior unicus- 1842. Amphisorex lesuerii Duvernoy. Mag. Zool. pate teeth on each side are small and evenly d’Anat. Comp. et Paleontol., 1842. 25:33. aligned (the fifth is peg-like and diminutive). There Type from Wabash Valley, Indiana. is no post-mandibular canal in the dentary. 1827. Sorex personatus I. Geoffroy St.-Hilaire. The pelage of the masked shrew is brown- Mem. Mus. Nat. Hist. Paris, 15:122. Type from ish gray or dark brown, with paler underparts eastern United States. (grayish white, smoky gray, or buffy white). The 1942. Sorex cinereus lesuerii: Bole and Moulthrop. feet are brown, and the bicolored tail dark Sci. Publ. Cleveland Mus. Nat. Hist., 5:95. See brown above, paler below; in winter, the up- also Jackson 1961, for southern Wisconsin. per parts are more grayish. Molts April through 1925. Sorex cinereus cinereus: Jackson. J. Mamm., June, and winter fur is acquired in October 6:56. (Jackson 1961). The sexes seem similar in size. Some representative external measurements for adults from Outer Island of the Apos- Sorex cinereus cinereus Kerr tle Islands, Brick Creek, Clark Co., and six young from Portage County (with cranial mea- The name Sorex by Linnaeus means shrews, surements as well) are respectively as follows: and cinereus means gray. The name Cinere- 94.0 ± 11, 105.9 ± 4.6, 95.5 ± 3.8; 38.14 ous Shrew used by Jackson (1961) and others ± 1.6, 39.4 ± 2.6, 38.2 ± 3.7; 10.3 ± 1.0,

86 THE WILD MAMMALS OF WISCONSIN 12.8 ± 1.1, 10.8; wts. 4.63 ± 1.1 g, —, —, N = 6: greatest length of skull 16.7 ± 0.3, cranial breadth 7.48 ± 0.2; interorbital breadth 2.69 ± 0.11 mm. Twenty specimens of various ages from Portage County averaged 16.4 ± 0.2 (16.1-17.1), 7.5 ± 0.2 (7.1-7.8), and 2.7 ± 0.18 (2.5-3.2) mm. Dental Formula. The teeth of the masked shrew number 32; five unicuspate teeth are present. The formula is I 1/1, U 5/2, P 4 1/0, M 3/3. (I = incisor; U = unicusp or “unicuspid”). Geographic Distribution. Jackson (1961) had no specimens from any of the southwest- em counties, suggesting this species did not

Maps showing geographic distribution of Sorex cinereus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 87 occur in these prairie counties. The species nant on dates from April 24 to October 24. has been obtained from most of them, and Two lactating females were seen on October appears to range throughout the state. 8 and October 18. The young are born pink, Status. The masked shrew is abundant naked and blind, after a gestation period of about throughout most of the state, and decidedly 18 days. They are about 13 mm long and weigh beneficial in controlling insects. The masked 0.1-0.3 g (Blossom 1932, Kilham 1951). After shrew is important in many food chains (see 12 days, young weigh as much as 2.8 g each Mortality below), and it preys on injuri- and are haired out. The erupted teeth are not ous insect pests of crops and spruce trees. yet visible. Teeth begin to erupt between 9 and Habitats. This boreal species ranges south- 14 days. After 19 days young are weaned and ward and westward into the prairies, and is often ready to follow the parents about. Buckner taken in dense, wet prairies. Usually it inhabits (1970) observed parents feeding numerous frit- wet forests, swamps, bogs, and marshes. It oc- illary butterflies to their young, gathered together curs in many habitats (Getz, 1961d, Long 1974), in the shade of a log. The butterflies had been including some sandy, grassy and forested. sucking water from wet sand. The nest of the masked shrew is com- Shrews do not live long in nature, and posed of leaves and grasses, located near the the populations may be practically categorized surface of the ground in a cavity, under a log, as two age groups. These have been classed stump, or rock. The shape about three inch- as I and II, or young and old adults. The young- es in diameter, with the inner nest scarcely of-the-year usually have longer skulls, with the more than one-half inch in diameter. There braincase more arched. Their teeth are usu- seems to be a single entrance (Jackson 1961). ally less worn. The cingulum or circular ridge Foods. A variety of insects, insect larvae, around the base of the crown portion of the worms, spiders, flesh of mice and other shrews, bicusp in young animals “is oppressed to the seeds, and other vegetal material comprise the dentary” (sic, i.e., the premaxillary, Hoffmeis- foods of the masked shrew. Frequencies are inter 1989; Diersing 1980). With increasing age sects (65.3%), small vertebrates (7.1), centipedes and growth the exposed root increases in (6.8), worms (4.3), mollusks (1.4), sowbugs (1.2), length and pushes the cingulum away from vegetal (0.9), Arachnida (0.9), and undetermined the alveolus. The distance seems directly cor- (10.9) (Hamilton 1920). The mammals eaten related with age, at least in comparing one may in some cases be carrion. Whitaker and specimen with another from the same place. Mumford (1972) examined stomachs of 50 Mortality. Sometimes carnivores refuse to masked shrews in Indiana finding 26 kinds of eat masked shrews owing to their odor, which foods, mostly larval moths, butterflies, crickets, probably results from the rectal glands. Wea- beetles (adults and grubs), and leaf hoppers. sels, skunks, foxes, badgers, wolves, and bob- Reproduction. Young-of-the-year masked cats all kill shrews (Murie 1936). Occasionally shrews do not normally breed. Their second large frogs catch and eat these tiny shrews, as year is perhaps the only opportunity. Breed- do snakes and birds of prey (Gould et al. 1964). ing commences in March and lasts into Octo- Even a brown trout caught one in Schoolcraft ber. Usually one, but as many as three litters a County, Michigan (Baker 1983). A variety of year, is produced. Males exhibit narrow dor- fleas and ticks infest these shrews (Jackson solateral “flank” scent glands during breeding. 1961, Timm 1975). A comprehensive list of They also occur on the flanks of females but parasites was published by Whitaker (1974). are much smaller. There are six mammae. Six Home Range and Density. Jackson (1961) pregnant females from central and northern says the home range of the masked shrew does Wisconsin, and one from Upper Michigan, not exceed one-fourth acre (or 1200 square averaged 5.3 (2-9) litter size. They were preg- yards). Ozoga and Verme (1968) found Sep-

88 THE WILD MAMMALS OF WISCONSIN tember populations contained 40 percent Specimens examined. Total, 530. Ad- young-of-the-year. In Marquette County (Upper ams, Ashland, Bayfield, Burnett, Calumet, Michigan), Manville (1949) found four shrews Clark, Columbia, Dane, Door (Rock Island, per acre (10/ha) in northern cedar swamp, and 1. Pedants Road, north end Washington Is- three per acre (7.5/ha) in black spruce. In Baran- land on Lake Michigan, 2. Swenson Road, ga and Marquette counties, Haveman (1973) 1/2 mi. E Jesson’s Place, 3. Jessens Place found 11 per acre (27.5/ha) in spruce swamp, on Swenson Road 2. Washington Island, 1. nine per acre (22.5/ha) in bogs, and four per Detroit Island, 2. Bailey’s Harbor, 1), Dou- acre (10/ha) in spruce barrens. Anderson (1977) glas, Florence, Forest, Green, Grant, Iowa, found 93 per acre (240/ha) in hardwoods. Deep Jackson, Jefferson, Juneau, Kewaunee, frost kills small mammals, both shrews and their Langlade, Lincoln, Manitowoc, Marathon, prey, and the cessation of reproduction prohib- Marinette, Milwaukee, Monroe, Oconto, its replacement of shrew populations suffering Oneida, Ozaukee, Pepin, Portage, Price, mortality. Density in shrew populations falls Racine, Rock, Rusk, Sawyer, Sauk, St. Cro- throughout the winter. ix, Taylor, Trempealeau, Vilas, Washburn, Remark. Long and Affeldt-Gehring Washington, Waupaca, Wood counties. (1995) reported a partially albinistic masked Other records.— Ashland Co. Apostle shrew showing the Valais goat color pattern. Islands including Madeline, Oak, Outer, Rasp- This pattern of pigmentation of the fur in berry, Rocky, Sand, Stockton and York isles which the anterior half of the animal is (Kantak, 1981). normally pigmented, and the posterior is white, has been reported as developmentally possible in only medium-sized mam- Sorex arcticus Kerr mals, such as goats. This single record of Arctic Shrew the Valais pattern in such a tiny mammal may falsify the mathematical rule of Valais color 1792. Sorex arcticus Kerr. The animal kingdom. p. pattern pigmentation and size-dependence 206. Type from Fort Severn, Hudson Bay (Murray, 1989). Patterns that show bicolored 1837. Sorex richardsonii Bachman. J. Acad. Nat. patterns white at each end also were never to Sci. Philadelphia. This name also used by be found (but see the Blarina account be- Lapham (1853), Strong (1883), Snyder (1902), yond). To make a long mathematical story Hollister (1910), Jackson (1908), and Cory short, little mammals, such as shrews, were (1912). Type from Saskatchewan. thought to develop too fast for such a con- 1858. Sorex pachyurus Baird. Mammals. In Reports strained pattern of pigmentation to develop. Explor. Survey... Pacific Railway Route, Additional Natural History. Significant works 8(l):20. Type from Pembina, North Dakota. on masked shrews and other shrews are written A homonym of Sorex Pachyurus Kuster, by H.H.T. Jackson (1928), Junge and Hoffmann 1835, from Sardinia. (1981), and Findley and Yates (1991). Geographic variation. There is a single geographic race in Wisconsin. Jackson (1961) Sorex arcticus laricorum Jackson referred shrews from southeastern Wiscon- sin to S. c. lesuerii (Duvernoy), but Hoffmeis- 1952. Sorex arcticus laricorum Jackson. Proc. Biol. ter (1989) determined its boundary was much Soc. Washington, 38:127. Type from Elk Riv- farther southward, in Illinois. S. haydeni, er, Minnesota. which some regard as a distinct species, reportedly occurs together with S. c. cinereus The scientific name literally means arc- in Iowa and Minnesota. tic shrew. The adjectives “tricolored” or “sad-

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 89 dle-backed” shrew were descriptive names ap- diploid number is 29 in males and 28 in fe- plied in the past. Never in the original de- males (Kirkland and Schmidt, 1996). scription, his revision of the long-tailed shrews Standard external and some cranial mea- or his book on Wisconsin mammals did Jack- surements are included here from several lo- son ever discuss the meaning of his word la- calities in northern Wisconsin (Drummond, ricorum. The Latinized name may be loosely one Ad, F. and Medford 4 Yg and Ad M), translated as “a denizen of larches” (Ameri- respectively: Total length, 116; 114.25 (106- can tamaracks). This is a fine epithet, for tam- 119), tail 45; 42.0 (39-45), hind foot 14; 14.5 arack marshes are a preferred habitat. (12-17), weights 7.0 g; 8.8 (7.8-10) g; great- Description. Arctic shrews are medium- est length of skull 21.1; 19.8 (19.6-19.9); sized, with long snout and long bicolored tail. cranial breadth 9.54; 9.1 ( 9.08-9.11); and The body and skull are larger than those in breadth across the maxillary processes 7.0; Sorex cinereus (see Measurements). Pelage in 5.01 (4.93-5.09). From Portage County, 16 summer, and in young-of-the-year, is brown Yg and Ad males measured 111.7 ± 0.74, dorsally, slightly paler below; in adults there is 40.5 ± 5.3, 13.5 ± 1.2,and N = 5 M, age a black mid-dorsal longitudinal band extend- class 2: 19.6, 9.47, and 5.05, respectively. ing from nape to base of tail, and the belly is Other measurements are listed in Kirkland and grayish brown. This pattern is tricolored: black, Schmidt (1996) and in Clough (1963). brown, and paler grayish brown. Three speci- Dental Formula. The dental formula for mens in molt were taken from late September Arctic shrews is the same as in Sorex ci- to late October. The cranium is arched, never nereus, but teeth are larger. I 1/1, U 5/2, flattened, the unicuspate teeth brightly pigment- P4 1/0, M 3/3 = 32. ed on the tips of unworn cusps, the third uni- Geographic Range. Common in Upper cusp tooth is larger than the fourth, the fifth a Michigan, but restricted from the Lower Pen- minute peg. The skull of the water shrew has insula by Lake Michigan, the Arctic shrew the third unicusp smaller. The rostrum, though ranges throughout most of the northern coun- narrow anteriorly, is broader across the cheek ties of Wisconsin, and occurs in southern Dane teeth. A post-mandibular canal is found in the and Dodge counties (Clough 1960; Lowell dentary in S. arcticus. Getz, personal communication). Arctic shrews have trivalent sex chromo- Status. The Arctic shrew is seldom com- somes, X and two Y’s. Females are XX. The mon, but lives in remote marshes and wet prairies so that it maintains fair abundance, except where these are drained or polluted. It has little economical importance, except that it feeds on larch sawflies and other forest insect pests. Habitat. Arctic shrews dwell in marshes, the grassy edge of cedar, spruce and tamarack swamps and bogs, stream valleys, and near some lakes. Whitaker and Pascal (1971) found them in old fields in Minnesota. This species seems limited to the boreal, coniferous forest region in North America, in the appropriate marshy and tamarack habitats. In central Wisconsin, I have taken them in dense grassy marsh, quite hummocky, but also where the black soil is level and evident Skull of Sorex arcticus. through the grasses. Cedar, alder, tamarack,

90 THE WILD MAMMALS OF WISCONSIN and a few spruce were at the edge of the marsh. Clough (1963) also caught them in hummocky marshes. Schmidt (1931) caught one in a sphagnum bog. Little information is available on homes of Arctic shrews. Reportedly the nests are globular “surface” structures comprised of grasses and other vegetation (Clough 1963, Baker 1983). Foods. Clark (1972) found some arthropods in several stomachs of Arctic shrews. Arctic shrews are often snap-trapped with rolled oats and peanut butter. In 62 specimens from Manitoba, the diet was exclusively

Maps showing geographic distribution of Sorex arcticus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 91 on snow. A long-tailed weasel killed one. There are various parasites such as fleas, mites, and ticks reported by Timm (1975), Kirkland and Schmidt (1986), and Whitaker and Pascal (1971). This shrew lives up to 18 months. Home Range and Density. There is no information on home range in Wisconsin Arctic shrew. P. Kim Van. 1995. Arctic shrews. They may occur in densities as high as five per acre (12.5/ha) (Banfield insects (Buckner, 1964), especially notewor- l974). Clough (1963) estimated densities as thy the pest larch sawfly. A captive from 3.5 per acre (8.8/ha). Haveman (1973) Stevens Point fed regularly on mouse carcass- found densities to be four per acre (10/ha). es, never feeding on dead songbirds. Bucker (1966) reported home range based Reproduction. Clough (1963), Baird et al. on 79 shrews to be 1.46 acres plus or minus (1983), and Baker (1983) reported litters as large 0.14 acres (0.6 ha). as ten, averaging seven. Birth occurs in April Kent (1983) found in his studies in Clark and May, and occasionally later. Jackson (1961) County, Wisconsin, that Sorex arcticus oc- suggested that September breeding indicated curred in Northern Hardwoods and Northern that a female might have two litters each sum- Sedge Meadow. He only trapped approxi- mer. Buckner (1966) found a few female young- mately 6 specimens in 1979-1980. Although of-the-year may breed. This suggests a typical Sorex cinereus and Sorex (Microsorex) hoyi pattern of small mammal breeding in Wiscon- were much more abundant, Arctic shrews sin. Arctic shrews have winter carry-overs, which were taken along drift fences at the rate of reproduce and may die, and young of the year 0.01 (N=3) and 0.02 (N = 3) per day in these reproducing later in summer. two communities. Whereas Kent found S. Nora Lopez-Rivera found an entry in the arcticus rare in these communities, Sasse Hamerstrom’s field journals of a pregnant (1978) found it the most abundant of any of Sorex arcticus taken 30 April 1979 in the the six shrews present in this same study area, Buena Vista Marsh. It contained 7 embryos. I except rare in the Hardwood Forest. collected a lactating female 1 mi. E. Jordan In 1976 (6-17 August), Sasse (1978) Pond on 5 August 1968, and Carl Becker caught up to 22 specimens per night in Marsh caught one 30 October in winter pelage with and Shrub communities. The Arctic shrew was teats barely visible. Embryo counts range from the most abundant mammal trapped in Shrub 4-9 in central Wisconsin ( mean 6.6, N=10). and Marsh communities, exceeding even the In Minnesota, near Minneapolis, 113 prolific meadow vole. I found Arctic shrews female Arctic shrews showed evidence of abundant in the Jordan Marsh, Portage Coun- breeding from April (lactation) into Septem- ty, co-existing with Sorex cinereus. ber (observed lactation). Reproduction ceased Remark. Clough (1963) observed that cap- by October. Males were reproductively active tive Arctic shrews were docile, never bit, and in from April to August. Young entered the pop- nature they are active either night or day. ulation in June. Most did not breed the first Geographic variation. There is no geograph- year. Litter size in Minnesota was 7.7+ 0.42 ic variation evident in Wisconsin and Upper Mich- SE, range 5-9 (N=10). igan. The shrews exhibit constancy in cranial and Mortality. Nelson (1934) found the re- external characters over a wide range. mains of an Arctic shrew in a great-horned Specimens examined. Total, 151. Bay- owl pellet. One dead Arctic shrew observed in field, Burnett, Calumet, Clark, Juneau, central Wisconsin was dropped by a coyote Langlade, Lincoln, Marinette, Marathon,

92 THE WILD MAMMALS OF WISCONSIN Portage, Price, Sheboygan, Taylor, Waush- standing water in a marsh, hence its vernacu- ara, Wood counties. lar name “Water Shrew.” Other records. Hamerstroms’ Journals, Description. The water shrew is large for Portage Co.: Buena Vista marsh. Clough, its family, largest of the Sorex in Wisconsin 1963. Jackson, 1961. (see Table Ins-3), with a long tail, long pointed nose, hind foot broad and fringed with stiff hairs as an adaptation to swimming. The cra- Sorex palustris Richardson* nium is large, broad and flattened; rostrum Water Shrew narrow but abruptly becoming broader span- ning across P 4/ to P4/ or M 1/1. The third “Every individual of this species [Sorex palustris] unicusp is smaller than fourth. was either in or at the edge of the water. In the The pelage, especially of the dorsum, of water they swam beneath the surface; encounter- the water shrew is slate gray to black, and ing obstructions... they crawled up and over usually blackish. Underparts of head, body and these.... one was seen June 30, 1924, making its tail silvery gray to brownish gray. There is an way up a small stream... it traveled on the ground, iridescent purplish sheen on the dorsum in but it took to the water where the bank became bright light (Baker 1983). Molt is observed too steep. Within a few minutes another individual on specimens in July-late August and early came along, following the path of the first.... January-March, suggesting two molts per year Along Manzanita Creek, June 6, 1926, a female (Jackson, 1928). was trapped at the opening of a small tunnel at the External measurements of seven Sorex edge of the stream. This tunnel except for its palustris adults from Seney Nat. Wildlife Ref- smaller size appeared like the ordinary surface uge, Upper Michigan, and USNM specimens tunnel of a mole. Active individuals were observed from St. Germain, Vilas Co., 1; Basswood at various times of the day, and on at least five L., Iron River, 1. Danbury, 2. Mercer, 1. Total occasions; twice animals swimming after dark were length, 148.3 ± 5.2 (142-156), length of tail, seen by aid of light reflected from a camp fire.... 65.7 ± 3.6 (62-73); 19.25 ± 0.89 (18-20). the animals appeared silvery, due to the air The sample size was N = 9. One adult weighed [bubbles] contained in their fur .” — Joseph 10.3 g. From several localities in northern Grinnell, Joseph Dixon, and J. Linsdale. In Wisconsin, 3 adult males and 7 adult females Vertebrate Natural History of a Section of averaged, respectively, as follows: 152, 150 northern California through the Lassen Peak ± 5; 76, 67 ± 1.4; 21.7, 18.57 ± 0.8; —, Region. Univ. California Publs., Berkeley. 11.91 ± 0.95 g, and five adult females aav- eraged in cranial measurements 21.42 ± 0.39; 10.2 ± 0.46; and 5.9 ± 0.21 mm. Sorex palustris hydrobadistes Jackson Dental Formula. The dental formula is the same in Sorex palustris as in Sorex cinereus. 1926. Sorex palustris hydrobadistes Jackson. J. DF = I 1/1, U 5/2, P4 1/0, M 3/3 = 32. Mamm., 7:57. Type from Withee, Clark Co., Geographic range. Occurs throughout the Wisconsin. Upper Peninsula of Michigan ranging southward in Wisconsin as far as Portage County. The name Sorex palustris means marsh Status. An interesting and poorly known shrew. Very seldom does this shrew range boreal species, the water shrew is chiefly in- away more than 2 m from a stream, lake or sectivorous and, if not beneficial, certainly harmless to humankind. In Portage County the * The nominate race’s type was from “marshes” [of shrew has seemingly disappeared from some Canada]. former habitats (e.g., Hay Meadow Creek), I

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 93 rola, baneberry Actaea rubra, bearberry Arctosphylos, and water cress Rorippa. The nearby forest was fir. In Canada I have taken the water shrew from the stony beach of the rushing outlet of a lake, with only a few grasses growing there, and in central Wisconsin from the black muddy sand along shallow creeks in marshy and swampy places, where the grasses and jewelweed were dense and tall. Little information is available on the burrows of water shrews. One nest was found in Skull of Sorex palustris. a beaver lodge (Seigler, 1956). The nest is about four inches in diameter, made of twigs suspect from the effects of insecticides. Possi- and leaves. Bankside burrows are used for bly the use of toxins, including fish manage- security, probably as homes (see quote pre- ment poisons, may adversely affect these semi- ceding this account). aquatic forms. Seldom abundant in any place, Foods. The water shrew feeds chiefly on the small numbers in museums suggest rarity. invertebrates, aquatic insects, water spiders, Jackson (1961) observed only 10 specimens snails, small fishes, and on occasion vegetal in all the collections he studied. Our collection material (Hamilton 1930, Whitaker and contains only 22 Wisconsin specimens. Lack- Schmeltz 1973, Buckner and Ray 1968, T. ing evidence to the contrary, the species may Clark, personal comm.). Buckner and Ray be regarded as in peril. Management of streams (1968) found carabid beetles the chief prey, and wetlands, therefore, should include this with larvae of Lepidoptera and Hymenoptera, species for protection. snails and other insects. In Montana, Con- Habitat. The water shrew dwells at the away mentions insects in 49 per cent of the edge of streams, bogs, lakes, rivers, and stand- stomachs examined, and the chief foods were ing water in marshes and swamps. It favors insects, a few planarians, and a little vegeta- rocky, log-strewn cold-water streams, with tion. Three shrews had fishes, either scales in numerous crevices and overhanging banks, the stomach or remains in the mouth. His and bordered by boreal woodlands (Borell and captive water shrews also caught tiny fish. Ellis 1934, Conaway 1952). This animal for- Occasionally he found mouse hair in a stom- ages on the banks, swims in the water, walks ach, one contained hair of a nearby eviscer- on the bottom, and runs across the water ten- ated water shrew. sion of still water (aided by light weight, large Reproduction. Very little information is feet and bubbles in the fur). Jackson (1961) known for water shrews in Wisconsin. The reports four water shrews from sphagnum and breeding season extends usually from January swamp laurel (Kalmia glauca) accessible to through August. Jackson (1961) suggests the water near Rhinelander. gestation period is 21 days. Several litters, usu- In western Montana, from the distant ally of six offspring (3-8) may be produced in a race S. p. navigator, Conaway (1952) made season. Young appear in the population by July, a fine study on this poorly known species. and probably breed their second year. The life The habitats he observed were always near span may not extend beyond 18 months (Con- cold-water streams with rapid currents, run- away 1952). A specimen taken near Stone ning through boulders, crevices, and stony Lake, in Sawyer County, had 3 embryos mea- areas. The vegetation along the water includ- suring 5 mm in crown-rump length on 9 April ed liverworts, mosses, green and sidebell Pa- 1969. Spiegler (1956) found four immature

94 THE WILD MAMMALS OF WISCONSIN water shrews in a water shrew’s home in the sticks of a beaver lodge (May 1954). They weighed 6.0-6.9 g (mean 6.4), and measured 72-75 mm in length (tail 24-27 mm). Conaway (1952) reported and reviewed reproductive records of water shrews, finding that in Montana the males begin sper- matogenesis in January and February. Fe- males show ovulation and lactation in March and as late as August. Three young females were breeding in June, and one was pregnant. No young males were observed to be sexually active. Old males were usually show-

Maps showing geographic distribution of Sorex palustris in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 95 ing sperm development, or at least some Subgenus Microsorex stages of gametogenesis, after the winter. In Pygmy Shrews Montana the streams were usually open in winter, and the shrews were active under ice The genus Microsorex was relegated to the shelves, but seldom on the snow. Being ac- genus Sorex and given subgeneric status by tive, perhaps reproduction was not impeded Diersing (1980). Some students (e.g., Junge much by winter. In any case, some embryos and Hoffman 1981, and others) have followed appeared in February. Altogether, Con- him in this arrangement, and others (e.g., Hall away’s records of litter size ranged from 5 1981) have not. Long (1972a) mentioned the to 8, with a mode of 6, and one female utility of the name, meaning in Latin “little showed the transfer of blastocysts to the op- shrew” or shrew-mouse (versus “shrew” or posite uterine horn (5: 1 to 3: 3). shrew mouse). That is rather appropriate for Conaway (1952) found only two age possibly the smallest terrestrial mammal on classes in water shrews. His age classes were earth. Do the pygmy shrews belong to a sep- based on toothwear studies, and related to arate genus or under the umbrella of Sorex reproductive maturity. spp? The answer is not straightforward. Mortality. Jackson (1961) listed preda- Diersing (1980) considered the Microsor- tors that occasionally kill water shrews as ex as one end of a transition, being an ex- weasels, garter snakes, and possibly hawks, treme both in diminutive third unicuspate owls, mink and large fishes, including trout teeth, and in hypertrophy of an inner “tine” (Doutt et al., 1966). Fleas, ticks, and mites or style along each upper incisor for about are reported by Timm (1975) and Whitaker half its exposed length. The tines are often and Schmeltz (1973). Conaway (1952) found worn away in older shrews (e.g., UWSP- two fleas, Neactopsylla and Corrodopsylla, 5014). He might have added another special- two mites Hirstionyssus and Euphagamog- ization, on the two anteriormost unicusps, amasus, 4 species of nematodes including which is the serial development of tiny but Capillaria, and two tapeworms Hymenole- distinct internal cuspules on the so-called “uni- pis and Tetrahyridium. cuspate” or “unicuspid” teeth, set off from Home range and Density. Little informa- the apices of these teeth where the ridge from tion is available for Wisconsin water shrews. the apex meets the cingulum. This trait is Buckner and Ray (1968) retrapped a shrew developed also in Blarina. in Ontario six times, from five traps, provid- This same evidence can be taken instead ing an estimate of home range as 0.8 acres. as evidence for generic status for these shrews, Another was captured four times, providing an estimate of 0.5 acres. The first was taken in August, the latter in September. Geographic Variation. None was observed in Wisconsin. Specimens examined. Total 27. Bay- field, Clark, Forest, Langlade, Lincoln Marinette, Portage, Sawyer, Taylor, Vil- as counties. Jackson’s (1961) records were examined, and also were mapped for comparison of range then and now. Vilas Co.: Lake St. Germain 1. Bayfield Co: Barswood Lake, 1. Sorex (Microsorex) hoyi. Philip Wright. Courtesy of J. Burnett Co. Paradiso & the Johns Hopkins Press.

96 THE WILD MAMMALS OF WISCONSIN for in these trenchant characters Microsorex possibly resulting from injury. The disk is a is not intermediate (Table Ins-3). Some shrews specialization, altered beyond the ancestral with a diminutive third unicuspate tooth, such typical unicuspate tooth. In company with the as Sorex palustris (in Otisorex) and Sorex other advancements, Microsorex seems dif- merriami (in Sorex), essentially lack tines on ferentiated from other Sorex. The homolog the bicusps. Repenning (1967) and Guilday (or analog) in Blarina (see below) suggests the (1962) mention another character, reduced origin of discoid form. entoconid height relative to the hypoconid, distinguishing ancient fossils of pygmy shrews from Sorex. Sorex (Microsorex) hoyi Baird There is no hybridization between Sor- Pygmy Shrew ex (Microsorex) hoyi and other Sorex, although opportunity exists, and indeed the two 1858. Sorex hoyi Baird. Mammals, in Reports Ex- kinds have been separated since the mid-Pleis- plor. Surv. for a Pacific Railway Route. Vol. tocene (Hibbard 1944). The crux perhaps is 8(partl):32. For 1857. Type from Racine, Ra- the importance of the peculiar third unicus- cine Co., Wisconsin. Also, Diersing (1980), pate tooth. Is it really the extreme form of a Long (1999a). transition? Or are diminutive third unicuspate 1858. Sorex thompsoni Baird. Mammals, in reports teeth in other soricine shrews a parallel in Explor. Surv. for a Pacific Railway Route. evolution? They retain pigmentation seen se- 8(l):34. See Long (1972a). Type from Burling- rially in the teeth, whereas in pigmy shrews ton, Vermont. the disk has no pigment, no inner loph, no 1925. Microsorex hoyi intervectus Jackson. Proc. function at all except, perhaps, as a spacer. Biol. Soc. Washington, 38:125. Type from In the evolution of crowded teeth and excep- Lakeland, Oconto Co., Wisconsin. tionally tiny body form spacing seems adap- 1901. [Microsorex] hoyi: Elliot. Field Columb. Mus. tive, and the “tines” of the upper incisors serve Publ. 4 Zool. Series, 2:377. the same use. The disk tooth seems functional, not vestigial, and is seldom lacking, ex- Sorex means shrew, and the name Mi- cept in an observed specimen (UW-SP 648). crosorex means little shrew. Philemus R. Hoy In this abnormal specimen the disk is lacking was a well-known amateur scientist from Ra- along with the adjacent tooth adjoining ante- cine, who sent a specimen to Dr. Fullerton riorly, on one side only, and this anomaly may Baird, who subsequently named the species have resulted from an injury. Feldhamer and in Hoy’s honor. Concerning “pigmy” versus Stober (1993) also reported a missing tooth “pygmy,” Hall’s students, except in this case, always went with American usage instead of a British or Canadian spelling. Jackson (1961) Table Ins-3. Characters of Wisconsin shrews: Blarina, and (1972) also followed that convention, pig- Sorex, and Sorex (Microsorex) hoyi. my over pygmy, but Hall himself used “pygmy,” and others followed suit. An appropri- Species “Tines” 4th < 3rd Cuspules Hypoconid Inner high ate French saying is, Je crois il va du blanc au noir, il est petit n’est ce pas? Blarina None No* Large Yes Description. Sorex (Microsorex) hoyii is S. arcticus Small Yes Small No S. cinereus Small Yes Medium No a tiny shrew with long tail, a pointed but rel- S. palustris Minute No Medium No atively broad snout, tiny eyes and feet, and S. (M.) hoyi Large No** Medium-large Yes five toes on each foot. It very closely resem- *U3/ is pigmented but approaching discoid form; bles the masked shrew in size, form and col- **U3/ is an unpigmented disk. or, but is slightly grayer dorsally, tail shorter,

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 97 and muzzle or snout less slender. The skull is grayer, sometimes whitish or buffy, and of- more flattened, especially in older shrews, with ten tinged about the throat with rusty brown. rostrum broader, cheek teeth with entoconid Molt occurs in late April-May and in October- reduced, compared to the hypoconid (poste- November (Long 1974). rior margin of the molar). The most trenchant Wisconsin pygmy shrews do not belong character is the third unicuspate tooth, which to the dwarf races of the subgenus Microsor- is minute and hardly visible, squeezed between ex (Long 1972; Whitaker and Mumford adjoining teeth as a wedge or disk, somewhat 1972), but the Racine specimens were ex- tilted and unpigmented. Difficult to observe, tremely small. One winter specimen, found even with a dissecting microscope, the disk is by Dr. Robert Freckmann, in Stevens Point, practically invisible to the naked eye. The tiny may be the smallest adult, terrestrial mam- fifth unicuspate tooth is also small and practi- mal ever found. Weighing only 1.8 g, and cally hidden by the tooth posterior to it, so having teeth extremely worn (indicating great that it is often said that when one casually age), the tiny specimen may have lost some examines the unicusp row only three teeth weight by desiccation, although it was found can be seen. In Sorex cinereus four (all ex- frozen in ice. Measurements and weights are cept the fifth) are visible and they are rather as follows for four adult males from Bayfield, even in form. In the pygmy shrew, the first four from Drummond, and one from Pray: and second unicuspate teeth are even, and Total length, 81.25 (80-84), 85.5 (84-88), conspicuous, the third cannot be seen, the 94; length of tail 29.5 (28-31), 29.8 (28-31), fourth is much smaller than the anterior two, 31; hind foot 9.75 (9-10), 9.75 (9-10), 9; 5.4 and larger than the fifth. The patterns can be g (N = 1), —, 2.9 g; greatest length 15.6, — suggested by the following: cinereus ////o , 15.4; interorbital breadth 2.8, —, 2.9; cra- and in Microsorex ll.lo where “o” is the tiny nial breadth 6.55, —, 6.6 mm. Adults weight fifth unicuspate tooth and the period is the about 4-4.5 g, in Wisconsin, but from the disk. This pattern can be discerned with care- same race Jackson (1928) reported 2.1 and ful examination with the naked eye. Addition- 2.9 g, from Elk River, Minnesota. ally the gap between the upper bicusps is filled, Dental Formula. I 1/1, U 5/2, P 4/ 1/ except in some old specimens, with inner 0, M 3/3 = 32. The disk is a permanent tooth. extensions called ‘tines.’ These spacers are Geographic Range. The pygmy shrew elongate, extending along and adjoined to probably is restricted from hot, dry prairies in each inner bicusp. the west and south, where few specimens have The pygmy shrew is brown or brownish been found. Specimens formerly referred to gray above with bicolored tail, underparts M. thompsoni (Long 1972a) were referred to S. hoyi hoyi (Diersing 1980), and in this I do not demur. Status. In many parts of the range of pygmy shrews the animals are spottily distributed, low in density, and over most of the vast range extremely rare. Some workers have taken them repeatedly with Sorex cinereus, but in the low ratios of say 50:1, or even 100:1. In southern Wisconsin the pygmy shrew is even rarer; only a few specimens have ever been caught. Their habitats are disturbed by man so much the species seems likely ex- Skull of Sorex hoyi. tirpated there. In central and northern Wis-

98 THE WILD MAMMALS OF WISCONSIN consin the shrew is uncommon but habitats for the species are being devastated by “urban sprawl”. Habitat. This boreal shrew inhabits most northern habitats (except the frozen tundra), occurring in a variety of climax forest and even seral habitats. Not found in dry prairies, it does occasionally occur in dry grassy areas, in tamarack, cedar, alder, and spruce swamps, with sedge grasses or sphagnum, in marshes, grassy meadows (dry, wet, even mowed), on stream banks, in coniferous and mixed coniferous forests, and even dry woodlands not too far from water. Beech-maple

Maps showing geographic distribution of Sorex hoyi in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 99 forest, pine barrens, and sphagnum bogs are in Wisconsin that would seem infrequent. all inhabited on occasion by pygmy shrews. Possibly obscuring the problem, many females Usually the habitat is near a stream or stand- caught may be young-of-the-year, and too ing water, and soils are patchily wet and dry young to breed. (Long 1972c). The use of pitfall traps pro- Feldhamer et al. (1993) examined tooth- vided a precious number of modern-day spec- wear to age specimens of Sorex hoyi, consid- imens to be accumulated. ered comparable to shrews of known age of In Manitoba, the pygmy shrew displayed other species. The age was subtracted from a wide tolerance for diverse habitats, except the date of capture to estimate birth date. They tundra, arid grasslands, and cultivated fields. found southern pygmy shrews breeding in all They were collected in fen, marsh, bog, and months of the year with a peak of breeding in shrub thickets, also deciduous and coniferous late winter. Using their method (but excluding forest and savanna (Wrigley et al., 1979). One old shrews), I estimate in Wisconsin the pyg- was taken in “prairie.” my shrews breed from June to September, No information is available on dens of occasionally later, with peak reproduction in the pygmy shrew. It is said to dwell in bur- July. Most females I examined were of young- rows, under a rock, or in roots of fallen trees. of-the-year shrews. In lower Michigan, none A captive shrew made several nests of cot- of 18 young-of-the-year pygmy shrews (Au- ton, open at both ends. gust 19-23) were sexually active (Baker, 1983). Foods. Little information is available. Mortality. House Cats catch pygmy Captive pygmy shrews have fed on flesh of shrews but seldom eat them because of the mice, other shrews, earthworms, but more rank odors from anal and flank glands. Other likely in the wild they feed on small insects predators include brook trout, garter snake, and insect larvae. The tiny, southernmost race and hawks (Cahn 1937, Long 1974). Fleas, S. hoyi winnemana fed on insect larvae, spi- mites, a tick (Ixodes), and hymenolepidid tape- ders, beetles, and ants (63 stomachs) accord- worms parasitize pygmy shrews (Buckner and ing to Whitaker and Cudmore (1988). Have- Blasko 1969, Long 1974). man (1973) found flies, beetles, other insects, Home Range and Density. Long (1974) spiders and a little sphagnum moss in the diet caught four pygmy shrews in about half an of our race, observed in Upper Michigan. acre in central Wisconsin over a period of Reproduction. In central Wisconsin pyg- several weeks. my shrews, there is no evidence of breeding The collecting was terminated on behalf more than once in the year. The only preg- of the rare shrews. Wrigley et al. (1979) stud- nant female was taken rather late (July), and ied the habitats and abundance of Sorex hoyi most females were not pregnant at all. Dr. in Manitoba. The shrew was most abundant Philip Wright (personal correspondence) men- in aspen forest and black spruce forest, two tioned to me evidence that in Montana one taken from tree-line habitats. In central and female seemed to be in the process of pro- northern Wisconsin the pygmy shrew is lo- ducing a second litter while nursing an earlier cally more abundant than that in some plac- one. Pregnant females carrying 2-7 embryos es, even outnumbering masked shrews. have been reported from 8 June to as late as Geographic Variation. Jackson (1925) 3 August (Jackson 1961, Long 1976, Scott named pygmy shrews from northern Wiscon- 1939, and Osgood 1938). Hoffmeister (1989) sin as Microsorex hoyi intervectus. He used mentioned a Manitoba specimen lactating in the vernacular names “Northwestern Pigmy September. If breeding regularly commences Shrew” and “Intermediate Pigmy Shrew.” in June and continues into September, more Supposedly it had a broader and higher brain- than one litter per year may be possible, but case than the nominate race hoyi. Baird’s

100 THE WILD MAMMALS OF WISCONSIN excellent illustrations (1858, for 1857) of his The lower anteriormost antemolar tends to holotype for the nominate race reveal that be only bicuspidate instead of tetracuspidate, the description of the nominate race was mis- and dorsally the red-brown labial walls are construed; it agrees closely with that of shrews nearly straight. from northern Wisconsin. I (1972a) placed intervectus in the synonymy of Microsorex hoyi hoyi (= Sorex hoyi hoyi). Blarina brevicauda (Say) Specimens examined. Total 49. Adams, Northern Short-tailed Shrew Bayfield, Burnett, Clark, Jackson, Juneau, Manitowoc, Portage, Price, Sawyer, Win- 1823. Sorex brevicaudus Say. In Long. Account of nebago counties. an exped. Rocky Mts., 164. Type from near Blair, Nebraska. 1858. Blarina brevicauda: Baird. Mammals, in Re- Genus Blarina Gray ports Explor. Survey . Pacific Railway Route. Short-tailed Shrews 8(l):42.

The dentition in Blarina resembles that of The name Blarina, proposed by J. E. Sorex in many respects. Large size notwith- Gray in 1838, may refer to Blair, Nebraska, standing, the short-tailed shrews are rather near the type locality. The specific part of the typical shrews. The ear is a narrow slit hid- binomen literally means short tail. Because den by fur of the head, and in this trait as well of its extensive burrowing and large size the as having a relatively short tail Blarina re- mammal is often called the “mole shrew” or sembles Cryptotis. Doubtless related to its size confused with moles. The burrows may de- and strength, the skull of this semi-fossorial scend half a meter below the soil surface in predator is robust, broad, with prominent ridg- friable soil (George et al., 1986). es and sharp lateral processes on the brain- Description. The trenchant characters for case for muscle attachment. Such processes the short-tailed shrew are given for the genus are not developed on the papery thin and above. Thorn-like lateral processes on the cra- smooth skulls of smaller shrews. The fifth nium, one on each side, and large size of the unicuspate tooth in Blarina is tiny and hid- skull identify the skulls of Blarina from those den between the adjacent teeth on either side, of other shrews. Compared to other Blarina, but it is pigmented and pointed like the oth- the northern short-tailed shrew has a larger, ers. There is a crista extending from the apex stockier body, bigger feet, and a somewhat to the inner cingulum. On the other unicus- blunt or truncated nose. Blarina brevicauda pate teeth inner pronounced cuspules are has a karyotype of 2N=48 to 50 chromosomes present. It is, therefore, slightly inacurrate to refer to these complex upper teeth as unicusps or unicuspids. The third unicusp is smaller than the fourth, extremely compressed as a sulcate disk, i.e., concave posteriorly to accommodate an anterior ridge protruding from the fourth. In this specialization and several others (Table Ins-3), for one the remarkable cuspules on the unicusps, there is some resemblance to Microsorex. The dentary is stout and deep to accommodate exceptionally large molars. There is a post-mandibular foramen. Skull of Blarina brevicauda.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 101 (George et al. 1986). Females have six mam- food chains of many predators (George et al. mae. In Wisconsin, B. b. brevicauda is typi- 1986, Baker 1983). The animal is common, cally larger and paler than B. b. kirtlandi. but contrary to what is often said (e.g., see The color of Blarina is slate black or Jackson 1961), is not the most abundant grayish black, often heavily suffused with Wisconsin mammal. dull brown. The undersides are similar but Habitats. This ubiquitous shrew is found paler. Molt occurs at least once (in August with varying densities in most terrestrial hab- and July) and is reported to occur in early itats of Wisconsin. It especially prefers wood- winter (Jackson 1961). lands with open floors, thick leaf litter or well- Northern short-tailed shrews are about rotted humus, and an abundance of rotted logs four or five inches in length, and the tail is and stumps. Not only in dry woods (e.g., the approximately an inch. Weights vary to 30 g. summit of Mosquito Hill near Green Bay), this See Table Ins-4. (Cahn 1937, Long 1974). species flourishes in wet, black soil of cedar- Dental Formula. I 1/1, U 5/2, P 1/0, M tamarack-spruce swamps. 3/3 = 32. The nest is made of various plants, even Geographic Range. The Northern short- hair. The tunnels extend either way from the tailed shrew occurs throughout Wisconsin, nest and may be directed downward. One doubtless in every county. The nominate race tunneled under our garage door and stole piec- is restricted to the Mississippi and the St. Croix es of dog food from the nearby food pan, drainage basins and the Wisconsin River val- carrying the pieces one by one to a garden ley, excluding Dane County and localities hose, then dashing to the burrow entrance. It southward. would tug “mightily” and with surprising Status. The short-tailed shrew is decid- strength to pull food pieces from our fingers. edly beneficial to man. It also is important in Blarina caches foods in its burrow.

Table Ins-4. Populations of Blarina and Cryptotis. Cryptotis were from Wisconsin and Illinois.

Total l. Tail l. Hind Foot l. Greatest l. Zygomatic Br. Max. Cranial Skull breadth Processes Depth Blarina brevicauda kirtlandi from Portage Country Adult F 115.3±9.7 22.5±2.6 14.44±0.81 23.67±0.48 12.56±0.31 7.93±0.24 6.59±0.25 N=17 Adult M 108.4±7.25 19.63±4.7 10.56±2.07 23.57±0.58 12.28±0.35 7.89±0.28 6.52±0.18 N=10 Old Ad. F 108 25 14 23.9 12.8 7.4 6.55 N=3 – – – 23.7 12.5 8.6 6.45 101 22 14 – 12.3 7.6 6.10 Blarina brevicauda brevicauda from West, Wis., Mississippi Valley Adult F 118.09±9.2 23.48±2.6 14.65±1.2 24.27±0.5 12.77±0.35 8.04±0.28 6.63±0.26 N=33 Old Ad. F 116 23.8 13.75 23.7 12.75 8.03 112-120 22-25 13.14 23.6-22.9 12.6-12.9 7.75-8.35 Cryptotis parva #6506 Sauk Co. 63 16 9.7 – – – – F. Alc. Illinois 60-94 15-19 9-11 16.1 3.4 7.4 – N=4

102 THE WILD MAMMALS OF WISCONSIN One day we found it drowned in the dog’s water pan (Museum specimen no, 5946). Other nest descriptions for Blarina are provided by Shull (1907) and Hamilton (1929). Shull’s nests, ranging from 12-15 cm in diameter, were comprised of tree leaves, grasses, sedges, nettle, goldenrod, and other plants. Hamilton found nests under rotting logs. One nest was of elm leaves and another of maple leaves, 30 cm below the surface. It contained seven young. Foods. Blarina, especially the larger species, prey extensively on small mammals and even small snakes. The saliva is a powerful

Maps showing geographic distribution of Blarina brevicauda in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 103 digestive fluid for insects, and toxic to mice ring-billed gull, five kinds of owls, red-shoul- and other small vertebrates (Martin 1981, dered hawk, water snake, even fishes (brown Pearson 1942, Lawrence 1945, Jeuniaux trout, green sunfish, largemouth bass). The 1961, and Thomasi 1978). The bite may be great horned owl is a very important preda- considered weakly venomous to humans. tor feeding on this shrew in Wisconsin (Err- Jackson (1961) presents a fascinating account ington et al., 1940). Doubtless many shrews by President Theodore Roosevelt of a short- are killed in winter by deep and enduring frost tailed shrew killing a pine vole and a garter and consequent food shortage. Spring floods snake. Hamilton (1930) described an attack may also take a toll. on a meadow vole lasting 10 minutes. Fleas, mites, chiggers, and even bee- The diet is chiefly insects and other in- tles (Leptinus) are ectoparasites (Jameson vertebrates (crickets, moths and caterpillars, 1950, Moore 1949, Timm 1975, Whitak- beetles and grubs, spiders, millipedes and cen- er and Mumford 1972, Scharf and Stewart tipedes, sow bugs, snails, worms, and also 1980, and Wrenn 1974). A comprehensive seeds and other vegetal material (De Byle review of parasites is provided by George 1965, Eadie 1949, Hamilton 1930, Ingram et al. (1986). 1942). Some insects, such as the larch saw- Home Range and Density. Especially fly, are forest pests. in thickets and woodlands, throughout Wis- Reproduction. Two or three litters may consin, short-tailed shrew densities may be be produced by a mother Blarina in one grow- high, leading Burt (1946) and Jackson ing season (Timm 1975). Seldom does a shrew (1961) to assume the species is the most live over a few months. One lived in captivity abundant small mammal in the Midwest. My 27 months (Baker 1983). Young-of-the-year studies suggest that the meadow voles (Mi- may breed in the first year (as did Mus. No. crotus pennsylvanicus) is much more nu- 6062, June 15, 5 embryos), and after the long merous in meadows, weedy and grassy prai- winter only 6 percent of carry-overs live to ries, and marshes, especially in northern produce a second litter (Pearson, 1944). I have counties; and the white-footed mouse (Per- only two records of pregnancy (each 5 embs). omyscus leucopus) is much more abundant Hoffmeister (1989) only had six records for in the sandy and shrubby habitats of south- the southern race (mean number of embryos ern, western and central Wisconsin. Blari- 5, range 2-7). He suggested that students do na usually ranked second or third in rela- not carefully look for embryos. Possibly the tive abundance with these two species. Jack- females stick close to their burrows when preg- son and I have two independent tallies of nant and are seldom caught. Nowak and Parad- museum specimens from Wisconsin: Blari- iso (1983) report the litter size as usually 5-7 na. Jackson 393, Long (herein), 376; Sor- but it may number as high as 10. After a ges- ex cinereus 329, 398; Peromyscus leuco- tation of 21 to 22 days, naked helpless young pus 537, 1,029; and Microtus pennsylvan- are born measuring 22 mm and weighing about icus 567, 548. These ratios, accurate in 0.8 g. After 17 days the furred young move in April 1991, probably have not changed. A and out of the nest. They are weaned at 25 Museum Index (Numbers of specimens pre- days (Blus 1971). served) is skewed in favor of the less com- Mortality. Although an unpopular food mon because one does not prepare all the for carnivores owing to odors from the anus common specimens. Nevertheless, Blarina and flank glands, the short-tailed shrews are is abundant and widespread in Wisconsin. important in the food chains of many preda- On the Upper Peninsula of Michigan, tors, including weasels, badgers, foxes, fish- Manville (1949) found 0.3 to 3.0 short-tailed ers, striped skunks, bobcat and house cat, shrews per acre in May (0.75-75/ha), and 8.5-

104 THE WILD MAMMALS OF WISCONSIN 11.6 per (21-29/ha) acre in June, in Mar- Blarina brevicauda brevicauda (Say) quette County. Haveman (1973) found 19 per acre (47/ha) in hardwoods, 7 per acre in For synonomy of the nominate race, see Blarina brev- spruce swamp, and 5 per acre in bogs. Getz icauda (Say) above. (1961d) found moisture important for any high densities. Nowack (1991) reported the Range. Western counties in the proxi- density to vary from 3 to 30 per ha. The home mate Mississippi River drainage, and includ- range was 0.2-0.8 ha. ing the lower Wisconsin drainage. See Map. Remarks. Specimen No. 5171, from Measurements. See Table Ins-4. King, Waupaca Co., has a pelage basally light Jackson (1961) ascribed larger shrews plumbeous gray; distally the hairs become from western counties to the nominate race, even paler with a buffy, yet silvery-white suf- smaller shrews eastward to B. b. kirtlandi. fusion. There are some darker undertones He believed males were larger than females. showing in the fur of this nickle-colored shrew. Measurements are given in Table Ins-4. The feet and tail were pale grayish tan, and Specimens examined. Total, 81. Ash- the tip of the nose was black. A specimen land, Barron, Bayfield, Buffalo Co.: Cochrane (Mus. No. 7816) taken 30 Sept 1994 in 1. Burnett, Clark, Douglas, Grant, Iowa, Stevens Point seems of the Himalayan (see Jackson, Jefferson, La Crosse, Monroe, above) color pattern, as described in Sorex Price, Richland, Sawyer, St. Croix, Taylor, cinereus (see Long and Affeldt-Gehring, Trempeleau, Vernon, Washburn counties. 1995). The muzzle is gray all around and in- Other records. Ashland Co.: Apostle Is- cluding the eye. On either side gray is ex- lands including Madeline, Oak and Stockton pressed in small patches above the ear ca- isles (Kantak, 1981). nals. All four feet are gray. The tail is gray and around the base is a grayish tone. The soles of the feet and the lips and nose were Blarina brevicauda kirtlandi pink. Most of the body was pure white. Eight Bole and Moulthrop days later, Mr. Dominic Berna, who lives on the same (Stanley) Street, brought in anoth- 1942. Blarina brevicauda kirtlandi Bole and Moulth- er, so closely resembling the first that surely rop. Sci. Publ. Cleveland Mus. Nat. Hist., 5:99. the two were siblings. The distance between Type from Holden Arboretum, Kirtland Twsp, the captures was less than a quarter mile. The Lake Co. Ohio. first shrew was trapped in a kitchen, with one normal colored Blarina. The second was Range. Eastern counties outside the trapped in a tool shed. proximate Mississippi River drainage, but in- Short-tailed shrews do not seem to be cluding part of the Wisconsin River drain- cannibalistic on other living Blarina, at least age. See Map. the near neighbors, and often they are found Specimens examined. Total, 294. Ad- living together. Males probably search females ams, Ashland, Columbia, Dane, Dodge, out by smelling anal and body odors. The male Door, Fond du Lac, Forest, Iron, Juneau, has a distinct flank gland (with unknown func- Kenosha, Kewaunee, Langlade, Lincoln, tion) prominent in the breeding season. Ago- Manitowoc, Marathon, Marinette, Mar- nistic threats are territorial displays, and pul- quette, Menominee, Milwaukee, Oconto, sating calls of high frequency are emitted as Oneida, Outagamie, Portage, Racine, the animal searches about (unfortunately for Sheboygan, Vilas, Washington, Wauke- the shrews, it may be heard by owls). These sha, Waupaca, Waushara, Winnebago, regularly emitted calls may aid in orientation. Wood counties.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 105 Genus Cryptotis Pomel whitish or grayish, and the tail bicolored Least Shrews (brown above, paler below). Geographic Range. The three known This diminutive prairie shrew is characterized Wisconsin specimens of Cryptotis are from chiefly by its small size and exceptionally short three southern counties. It should occur in tail. The short row of unicuspate teeth has marshes and weedy prairies farther southward. only four, the posteriormost being a tiny peg- Status. The least shrew is possibly ex- like tooth practically hidden from view. Su- tirpated in Wisconsin. Always rare, no least perficially the antemolar teeth resemble those shrews have been obtained in Wisconsin of Microsorex, because the tiny, disk-like since 1944. This shrew should be reintro- tooth in the Microsorex is difficult to see. Both duced in the Wisconsin fauna, into preserved shrews show three of the teeth, and a poste- and restored prairies in southern and west- rior fourth is hidden, a small peg. The molars ern Wisconsin. The species, although one in Cryptotis have high crests in unworn teeth. of our rarest mammals, is decidedly benefi- On the upper unicusps there are prominent cial to humankind. cuspules, as in Blarina and to some extent in Habitat. The least shrew prefers grassy Microsorex. All the other shrews in Wiscon- prairie or grassy and weedy old fields, broom- sin have 32 teeth, but Cryptotis has only 30. sedge, briers, cultivated fields, grassy mead- The ear is hidden in the fur of the head. The ows, open woods and, rarely, orchards. lower anteriormost tooth is well developed but Herbaceous ground cover may be more never tetracuspidate to my knowledge. The important than soil type (Whitaker 1974). outer rim is nearly straight-sided. Little information is available on homes for least shrews. Jackson (1961) reports from other studies that the nest is globular, Cryptotis parva (Say) about four or five inches in diameter, com- Least Shrew posed of dry grass or leaves. There are usually two openings into the nest, which is 1823. Sorex parvus Say. In Long, Account... Rocky sited under a rock or log, and sometimes Mts., 1: 163. Type from Missouri River, Wash- underground at a depth of four or five inch- ington Co.: Nebraska. es. From Texas, Broadbooks (1952) de- 1842. Brachysorex harlani Duvernoy. Mag. de zool. scribed a nest found under a sheet of corru- de Anat. Comp. 25:40. Type from New Har- gated iron, about 100 m from a cat-tail mony, Indiana. marsh. About 7 inches in diameter, and 2 1858. B[larina]. eximius Baird. Mammals. Reports inches high, the nest was comprised of old Expl. for a Pacific Railway Route. 8:52. Type willow leaves. An opening from a short tun- from DeKalb Co., Illinois. 1912. Cryptotis parva: G. S. Miller, Jr. Bull. U.S. Nat. Mus., 79:24.

The name Cryptotis parva means a small animal having its ear hidden. Description. Same as given for the genus. From the only Wisconsin skin of Cryp- totis available, preserved in alcohol, the col- or is brownish gray. Illinois shrews are similarly colored. Some least shrews to the southwest have a chestnut dorsum. The belly is Skull of Cryptotis parva.

106 THE WILD MAMMALS OF WISCONSIN nel entered the nest from below and a runway connected to it from the side. There was a “toilet area” or latrine at the edge of the nest, where numerous small insects were feeding. This latrine was about 3 inches in diameter. If disturbed, the young, barely haired out, tried to hide in the tunnel or burrow into the leaves. Foods. Hoffmeister (1989) summarized foods in Illinois as lepidopterans, grasshoppers, crickets, chinch bugs, spiders, worms, and unidentified arthropods, and occasionally seeds and flower parts. Hamilton (1944) in

Maps showing geographic distribution of Cryptotis parva Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER INSECTIVORA 107 Indiana reported insects, earthworms, mol- Order CHIROPTERA lusks, centipedes, and some vegetation in di- BATS ets of least shrews from eastern states. Reproduction. In Illinois, young of Cryp- “If bats are to remain [on earth].... we must initiate totis parva are born in spring and fall, with conservation measures.... certain needs are obvious. the number of young varying from 4-7 Some caves harboring important colonies should be (Hoffmeister, 1989). Whitaker (1974) report- protected from undue human disturbance. The ed embryo counts as 4.9 (2-7) and litter size trend away from the use of the dangerous residual as 4.5 (2-7). Gestatation is about three weeks, pesticides must be continued. People who work with and the young are haired with opened eyes bats should try to minimize the disturbance.... in two weeks. Newborn weight about a third Finally, more people should become aware of the of a gram (Whitaker 1974). They are “full many aspects of a bat’s life; such awareness should grown” in a month. Conaway (1958) presents make people more interested in the protection of data on growth. these fascinating animals.” — Wayne H. Davis, in Broadbooks (1952) quotes Hamilton, Bats of America, by the late Roger W. Barbour that the young are born blind and naked and and Wayne H. Davis, 1969. grow 0.1 to 0.5 g per day for about 3 weeks. Broadbooks also found an adult male in at- The Order Chiroptera is comprised of tendance with the female at the nest. In cap- the only mammals that fly. Known popular- tivity, the female ate her young. Captive-pro- ly as bats, these aerial mammals are the only duced litters averaging 4.56 young (range 1- ones with wings, which are modified fore- 9), weaned in 18-19 days. These were sexu- limbs having elongated fingers and elastic ally mature in about 50 days. Probably least webs between them. Distal claws disappeared shrews breed from March until November, from the fingers of the hand, except on the although October young would find cold functional thumb [and on the index finger in nights and frost adverse. They live about 8 some primitive, tropical bats called flying months, but as long as 31 months. foxes]. Flight is gliding and thrusting the body Mortality. A red fox killed but did not eat through the air, with the capability to ele- a Wisconsin least shrew (Hanson, 1945). Great horned owls ate the other two Wiscon- sin shrews, subsequently found in their owl pellets. Hawks, owls, snakes and other shrews are likely predators. A flea and several species of mites parasitize least shrews. Home range and Density. Hoffmeister and Mohr (1957) reported populations in Illi- nois as high as 10-15 per acre, but in some years none. Geographic Variation. There is a single subspecies known from Wisconsin, and central Illinois. Specimens examined. Total, 1. Colum- bia Co.: Sauk Prairie, 1. Other Records. Total, 2. Jackson (1961): Columbia Co.: E of Prairie du Sac, 1 mi. E- SE Dam on Wisconsin River, 1. Dane Co.: Skeleton of bat. G. Cuvier, Le Règne animal. Woodcut. Pine Bluff, 1 mi. N Klevenville, 1. 1817, 1829.

108 THE WILD MAMMALS OF WISCONSIN vate trajectory. Volant locomotion, seen in lifted, although drag greatly increases. As the several other distantly related mammalian angle of attack is inclined the bat stalls. If one groups, such as flying squirrels, is simply glid- wing is extended into the airstream while the ing (“volant” actually means flying). Bats, other is held close to the body, the bat “flits” probably descended from primitive insecti- (Eisentraut 1936). vores. The molar teeth are formed similarly Many echolocating bats hawk insects. in W-shaped patterns. Ancient fossilized bats They rely on echos from high-pitched calls to (Jepsen, 1966; Simmons and Geisler, 1998), “see” in the dark with their ears (Griffin 1958, Icaronycteris index, Archaeonycteris, and others). Palaeochiropteryx, and Hasianycteris are There are two suborders of bats, the considered insectivorous, some with insects Megachiroptera, or flying foxes, in the Old in their stomachs, and they all had advanced World tropics and Australia, and the wide- wings. These wonderfully preserved fossils spread, abundant, and diverse Microchi- from the Eocene epoch give a few clues to roptera. The Microchiroptera have specialized the evolution of wings and flight. wings, in most cases tiny but functional eyes, In addition to toe webbing, there is in and have evolved several different patterns of bats a flight membrane on either side called a echolocation. Novacek (1985) reports that the patagium. It is a double-layered membrane of oldest known bats (from the Eocene of Wyo- skin extending laterally on each side out to ming and Germany) possess cranial features the wrist, to the tip of the fifth digit, and to suggesting an early function of echolocation. the ankle or hind foot. Such a lateral mem- Possibly the orientation sense arose in caver- brane has evolved repeatedly in arboreal mam- nicolous bats. mals, probably serving to hide the revealing Other features include the upright rod in shadow when the body is flattened against a the ear (the tragus), the palatal notch sepa- branch (Spurway 1955) or to break the fall as rating the premaxillaries (and their incisors), the animal leaps from a tree. A tail membrane, the soft rod attached to the ankle helping to found only in one other group (Order Der- make the uropatagium taut (called the calcar), moptera) stretches between the hind limbs (the and the flattened chest. uropatigium). Usually in bats the tail vertebrae are comprehended in this membrane. Several characters indicate a gliding origin of flight: bats and gliding mammals are all nocturnal, they fall into flight, bat patagia resemble gliders, and bats’ hind limbs could never leap from the ground. The glider has a great deal of potential energy accumulated by climbing (Long et al. 2003). When bats pull their wings downward and forward, by means of several pairs of relatively large shoulder and chest muscles, the forward edge of the wing is canted downward. Then air is thrust mostly behind, and the hind limbs join in this pumping thrust. Lift is achieved. The wind helps raise the wings to the power stroke position. When bats glide, aerodynamic lift is obtained from pitching the wings against the airstream. The light body is Big Brown Bats on Ceiling of Twin Bluffs Cave. 1977.

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 109 Table Chi-1. Incidence of rabies in bats of Michigan* Owing to a national interest in Lasionyc- (1965-78) and Illinois (1965-82). Of all bats tested, 5.8 per- teris noctivagans, and its particular strain of cent were rabid. rabies virus detected in humans, workers at MICHIGAN ILLINOIS the Wisconsin Rabies Laboratory must deter- Species Tested Rabid & % Tested Rabid mine which bats are dangerous and which are Myotis keenii 27 0 0 0 0 not. For example, an elderly man attacked by M. lucifugus 75 0 0 43 4 a bat in Texas, with no visible bite marks, L. noctivagans 12 2 16 90 4 contracted rabies and died a couple months E. fuscus 1,093 48 4 355 15 later (1997). He and three other Americans L. borealis 55 3 5 378 34 died from bat rabies that year, all attributed L. cinereus 12 2 16 134 20 N. humeralis 10 0 0 0to a virus of the silver-haired bat, Lasionyct- Totals 1,275 55 4 1,000 eris noctivagans. Two cases are of great interest. A recent *Figures slightly revised. Wisconsin death (November 1, 2000) of a man near Reedsburg, associated with silver haired bats by the viral strain, but health Rabies in Bats authorities blamed bats from his apartment. Those likely would not be silver-haired bats. One of only a few human deaths attributed Silver-haired bats migrate out of Wisconsin to rabies transmitted by the overt bite of a southward in October, and do so in large num- bat (species unknown) happened near Blue bers at this time (~ October 14). Mound, Wisconsin (Jackson 1961, Long September 4, 2004, a recovering 15-year old 1976). A man with no screen on his window patient, Jeanna Giese, relates catching a “small, black was bitten on the ear. He did not seek med- bat” [Lasionycteris is the only black bat in Wiscon- ical treatment used against rabies virus. He sin] in a Fond du Lac church. She treated a bite on contracted the disease. About 30 years ago her hand with peroxide, but developed advanced I read a report in the local paper by the Por- symptoms of rabies October 15, and was transferred tage County Health Officer that all bats were from St. Agnes to Children’s Hospital on October 18. found to harbor the rabies virus. What this Dr. Rodney Willoughby, infectious diseases and pedi- really meant was that the bats he sent to the atrics, and with the family’s support, attempted an State Hygiene Laboratory in Madison all har- untried treatment, using a drip treatment of the neu- bored the virus. After examining the reports, ral protective agent Ketamine, he was able to fine I suspected the number of bats he reported tune the effect of Ketamine with Benzodiazepine. After did not exceed four. In Wisconsin tabulations two days or so he added Ribavirin and even a flu drug (see Table Intro-2), few bats test positive. In Amantidine. The drugs and induced coma protected comparison, numbers of dogs, cats, and even the Central Nervous System. The immune system ful- cows (bitten by some rabid mammal most ly responded, and the patient progressed steadily. After likely) that were not inoculated against ra- about 80 days she was returned home, and has pro- bies showed higher incidence (but no longer gressed daily. Willoughby says enthusiastically that is this true since implementation of dog im- “She will go to college”. This is the premier triumph munization). Striped skunks (Mephitis me- over rabies. I reiterate, “Don’t pick up sick or wild phitis) showed a high, often epidemic inci- bats; tell your children to leave helpless bats alone. If dence. Constantine et al. (1979) found only one is bitten, take rabies shots.” 0.025 percent of naturally occurring bats In Michigan, Baker (1983) and Kurta harboring the virus. Other studies have (1979) reported of 1,247 bats examined by shown about one percent or less (Trimarchi the Michigan Department of Public Health only and Debbie, 1977). 65 (5 percent) had rabies, with only 3 to 11

110 THE WILD MAMMALS OF WISCONSIN positive bats in any years. In contrast, 28 per- tional, Austin, Texas, provides a handbook cent of 384 skunks were rabid. Both 5% and for building and siting bat houses. 28% are overestimates, because any animals captured while sick are more likely to be tested. In nature the percentages would be lower. Family VESPERTILIONIDAE Gray Trainer (1957) and Long (1976) called Evening Bats for a better study of the bats in Wisconsin. In 1991, I begain identifying bats with the State “The Ordinary Bats (Vespertilio, Cuv. and Geof.) — Laboratory of Hygiene (J. Powell). The UW- Have no leaf or other distinctive mark on the muzzle, SP collection was for four years a repository and the ears separated; four incisors above, of which (Tables Chi-2-4) for non-rabid Wisconsin bats. the middle two ones are apart, and six below, sharp- edged and somewhat notched: their tail is comprehended in the membrane.”— Baron Georges Cuvier, Bats in Houses in Le Règne Animal, 1817, 1846.

The Bat. It has no feathers, yet has wings, It’s quite This large and cosmopolitan family con- inaudible when it sings, It zigzags through the tains all the Wisconsin species. See accom- evening air And never lands on ladies’ hair. — panying figures, the key to Wisconsin bats, Ogden Nash. and the table of characters of Wisconsin bats. Vespertilionid bats in temperate and bore- In Britain, many people introduced them al latitudes, such as those in Wisconsin, show a into their attics where the guano is consid- complex interrelation of microhabitats (mi- ered insulation or collected for rich fertilizer. crotemperatures), migration, lactation, and Most Wisconsin people eradicate bats, but growth. Growth rates of young raised in mater- especially in older houses, bats may be found nity roosts vary with air temperature. In tree in any season, especially in autumn. dwelling bats, in their cooler roosts, the young There is a myth that sprinkling moth crys- and mothers often save energy by entering tor- tals will drive bats away, or stringing up an por. Their growth rates seem low. Bats contin- extension light will keep them out. Bats usu- ue growing throughout pre-fledging, pre-migra- ally leave a house at twilight emitting a few tion, and migration until the next season of re- drops of urine. Telltale stains on the eaves production. Lasiurus do not seem to lose body may mark the entrances, and tin or alumi- mass by lactation and tend to produce twins num must be used to cover the hole when the [even 3-4 young] regularly. They may forage bats or their young are not present in the primarily to feed the young, which do fledge at house. Small cracks may be caulked. The best an early (low) percentage of adult body mass way to remove bats is to put on leather gloves (61 percent in Lasiurus cinereus, 55 percent and grab them as they sleep. Sometimes it is in Eptesicus serotinus, which regularly has practically an impossible task to rid a house twins). The growth rates of the pre-fledged of bats. Near the Trees for Tomorrow camp young are lower than those in vespertilionid bats and nature center, in Lincoln County, is a with single young. In all Wisconsin species there man-made bat shelter that harbors many are two pectoral mammae, except Lasiurus Myotis lucifugus, an interesting place where (with four). In Lasiurus, with such a slow growth adults and children are taken to observe the rate even in a short growing season, by migra- evening flights. The house is sited about 30 tion to warm regions, little or no hibernation, feet high and houses. Many “bat roosts” for and feeding on available insects year round, the backyard are sold today. Most of them there seems to be compensation in maintain- seem ineffective. Bat Conservation Interna- ing energy budgets (Koehler and Barclay, 2000).

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 111 It is now possible to identify some bats Key To The Wisconsin Bats by their echolocation calls. Ultrasonic bat detectors today (see advertisements in the 1 Tail not extending beyond tail membrane Journal of Mammalogy) provide detailed (uropatagium) or extending less than 4 information on species identification of fly- mm beyond it ...... 2 ing bats, spatial and temporal bat activity, 1’ Tail extends distinctly beyond the uro- habitat use and prey selection, and even the patagium (known from northern Illinois, physiological processing of echoes (Fenton, but unverified for Wisconsin, a possible 1981; Miller and Andersen, 1984; Thomas wanderer)...... Mexican free-tailed Bat and West, 1984; Thomas, Bell and Fenton, Tadarida brasiliensis 1987; and others). This procedure promis- 2 A single pair of upper incisors ...... 3 es in this century to provide a superb sur- 2’ Two pairs of upper incisors ...... 5 vey of free-ranging bats in Wisconsin and 3 Upper surface of uropatagium densely the Upper Peninsula, whereas previously we furred, the skin not visible, two pairs of relied mostly on banding of cave species or upper premolars, one of which is a by netting to study the diversity and sea- minute peg, usually a whitish spot visible sonal population changes (see table Chi-5, on shoulder, dorsal pelage more or less & Fig.) on seasonal fluctuations and hoary ...... 4 abunance). 3’ Upper surface of uropatagium nearly na- Bats have an undeserved bad reputa- ked, skin visible and scantily haired prox- tion. For some inexplicable but deep emo- imally, one pair of upper premolars, tional feeling, or possibly (some would say shoulder spot lacking, dorsal pelage probably) it may be taught to people to some brown ...... Evening bat extent, there is an apprehension of bats that Nycticeius humeralis is often expressed by hatred and disgust. 4 Dorsal pelage maroon brown, washed Bats normally do not attempt to get into a with hoary whitish or buff, greatest length woman’s hair. Neither are they usually ra- of skull exceeds 16 mm, forearm great- bid. Bats are highly beneficial, limiting the er than 45 mm ...... Hoary bat populations of harmful insects during the Lasiurus cinereus “night shift” when most insectivorous birds 4’ Dorsal pelage brick red orange or yellow- are sleeping (E.R. Hall, personal communi- ish orange, thinly washed with hoary buff, cation; Barbour and Davis, 1969). Bats greatest length of skull less than 16 mm, should be protected, i.e., managed. Some forearm less than 45 mm ...... Red bat American bats are listed as Endangered Lasiurus borealis Species because they are so scarce and are 5 Dorsal pelage dark brownish or blackish vulnerable to destruction (in their nursing washed with buffy or silvery white ...... colonies or hibernacula). Kurta and Teram- ...... Silver-haired bat ino (1992) and others also showed the ad- Lasionycteris noctivagans verse effects of urbanization to bats in De- 5’ Dorsal pelage brownish, reddish or yel- troit, decreasing their abundance, species lowish buff, never washed whitish ..... 6 diversity and possibly their reproduction 6 Dorsal pelage yellowish, tan or pale red- success. In suburbs, with numerous shade dish brown, one large and one minute trees and attics, some species may increase. pair of upper premolars (total of 34 teeth Barbour and Davis (1969), Kunz (1988), and teeth), hairs conspicuously tricolored Nowak and Paradiso (1983) are important (brown/ cream/ gray), forearm usually references on American Chiroptera. less than 36 ...... Georgian pipistrelle Perimyotis subflavus

112 THE WILD MAMMALS OF WISCONSIN 6’ Dorsal pelage brown, or yellowish, pale, or reddish brown (faded pelage); normally one large and two minute pairs of upper premolars (total of 38 teeth) or only one large pair (total of 32 teeth); hairs never tricolored; forearm more than 36 mm...... 7 7 Skull more than 18 mm in greatest Vespertilionid bat. W. A. Weber, with Karl Schmidt. length, forearm more than 44 mm, premolars 1/2 (total of 32 teeth)...... Big brown bat or slightly keeled, sparse hairs extend be- Eptesicus fuscus yond toes ...... Little brown bat 7’ Skull less than 18 mm in greatest length, Myotis lucifugus forearm less than 40 mm, premolars nor- 9’ Dorsal pelage dull brown, dorsal and ven- mally 3/3 (total of 38 teeth)...... 8 tral pelage tinged pinkish or purplish gray 8 Ear pinna broad and long (16 mm or more over brown, calcar distinctly keeled, from notch), maxillary tooth row more sparse hairs usually do not extend be- than 5.5 mm, elastic fibers of uropatagium yond toes ...... Indiana Myotis forming a pattern of wide, prominent Myotis sodalis chevrons numbering approximately 7 (6- 11) ...... Keen’s or Northern Bat Myotis Myotis keenii To his adroit Creator ascribe no less the praise — 8’ Ear pinna narrow, more recurved, and Beneficient, believe me, his eccentricities. — shorter(usually less than 15 mm), maxil- Emily Dickinson, about 1876 lary tooth row less than 5.5 mm, elastic fibers of uropatagium forming narrow, General changes in abundance in Wis- indistinct chevrons numbering approxi- consin’s bats seem profound. Old literature, mately 12 or more ...... 9 including Jackson (1961), suggests the red bat 9 Dorsal pelage with metallic, brassy or was most abundant, the big brown bat un- coppery brown glint, never with pinkish common, but the situation is now reversed. or purplish suffusion, calcar not keeled Myotis lucifugus, ranging through forests and

Chart of Characters of Wisconsin Bats. Diagnosis to make comparisons easy. Asterisk means unknown in Wisconsin. P = small premolars.

Length Length Calcar No. Species forearm tragus keeled incisors Color Other P. subflavus 32-6 ~½ ear — 2 Yellow-brown Hair tricolor 1 P M. lucifugus 34-8 < ½ — 2 Brass-brown >18, elastic fibers M. keenii 34-8 ~½ ?+ 2 Brass-brown 7-12, elastic fibers M. sodalis 34-8 < ½ + 2 Dull-brown M. leibii* 30-34 < ½ + 2 Yellow-brown Black mask N. humeralis 34-8 < ½ — 1 Ashy-brown L. borealis 36-43 ~½ + 1 Whitish/reddish Round ears 1 peg P. L. cinereus 50-8 ~½ + 1 Whitish/brown Round ears 1 peg P. E. fuscus 44-8 < ½ + 2 Brass-brown Broad nose No tiny P. L. noctivagans 39-43 < ½ Short — 2 Whitish-black Tadarida brasiliensis* 40-5 no calcar 1 Gray Free-tail

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 113 Van Zyll de Jong (1985) believed that Myotis keenii found in the Pacific Northwest (and southwestern Canada) is specifically distinct from the wide-ranging eastern bats referred to the same species. Therefore, he elevated the eastern race M. k. septentrionalis to a full species, called M. septentrionalis. The bats are allopatric, and, therefore, the question whether they do or might hybridize is subjective, i.e., it is difficult to dem- onstrate. Lacking clear-cut evidence, one may emphasize either the resemblance of the two bats or their distinctness. In comparisons of Uropatagium of Myotis lucifugus (left) and M. keenii. some small samples, there was no overlap Note elastic fiber patterns and long ears. (Fitch and Shump 1979), but Miller and G. M. Allen (1928: 109-110) had clearly revealed and suburbia, was always abundant. Pipist- almost total overlap in lengths of body, tail, relles seem to be declining in numbers. Abun- skull, and maxillary toothrow, and breadths dance is analyzed in tables Chi 2-5. of zygomata and braincase. In 1994, I examined, with Robert Fish- er, several M. keenii from Alaska and south- Genus Myotis Kaup ward preserved in the U.S. National Museum Mouse-eared Bats (Nat. Hist.). We examined the uropatagia, having found myself an undescribed and subtle Small brown bats having fairly well-developed difference between keenii and other Myotis tragus, a mostly naked uropatagium, and P in the eastern states (see Fig.). Wide-spaced 1-3/1-3 present in Wisconsin species. and parallel elastic fibers in the uropatagium were also observed to be characteristic of the dark, coastal western specimens, as were the Myotis keenii Merriam elongate ear pinna and tragus. For the afore- Myotis keenii septentrionalis (Trouessart) mentioned resemblances and this new char- Keen’s Myotis or Northern Myotis acter, I retain the name M. keenii, reinforced by his astute judgment of Gerrit Miller, as well 1895. Vespertilio subulatus keenii Merriam. Amer. as the overlooked overlaps showing close re- Nat., 29: 860. Type from Massett, Graham Is- semblance. land, Queen Charlotte Islands, British Columbia. Description. The trenchant external 1897. [Vespertilio gryphus] var. septentrionalis characters of Myotis keenii are the large ear Trouessart. Cat. Mamm. . Fasc. 1, p. 131. Type conch with a correspondingly elongate point- from Halifax, Nova Scotia. ed tragus within, and the pattern of elastic 1928. Myotis keenii septentrionalis: Miller and fibers of the uropatagium widely spaced and Allen. Bull. U.S. Nat. Mus., 144:105. nearly parallel, like a music staff, and vary- 1985. Myotis septentrionalis Trouessart: Van Zyll de ing in number usually from 7-12 rows. In Jong, Handbook on Canadian Mammals, Part 2. Myotis lucifugus the number of fibers, crowded and confused, usually exceeds 18. The generic name means mouse-eared. Dr. In subadults and a very few peculiar speci- C. Hart Merriam named the species in honor mens the fibers are indistinct. In live or fresh of J. H. Keen, who collected the holotype. bats (only) one can push the ear pinnae for-

114 THE WILD MAMMALS OF WISCONSIN ward to see if they extend at least 4 mm be- The pelage of Myotis keenii has a bright yond the nose. This “book” character does sheen, which in bright light is called a “brassy not identify dry museum specimens. The glint.” The basic tone is brown or slightly yel- paraboloid shape of the elongate and wide lowish brown, but the sheen gives a golden pinna suffices to distinguish keenii from lu- tone to the brown fur. This pelage is usually cifugus, which has shorter, narrower and indistinguishable from Myotis lucifugus, Ep- somewhat recurved pinnae. One questions tesicus and Nycticeius. The patagia and ear whether the calcar is or is not keeled pinnae are slate or brownish gray. The ven- (Hoffmeister 1989, Baker 1983), for Wis- ter (or belly) is grayish or brownish buff, over- consin specimens are slightly keeled. lain with light buff, cream, or silvery gray. Molt The Myotis keenii skull is approximate- occurs in the summer. ly 15 mm long by 9 mm across the zygoma- Fitch and Shump (1979) provide mea- ta. The hind foot measures up to 10 mm, surements as follows: forearm length 36.4 occasionally 11 (see Measurements). The skull (35.9-37.0), ear length 18.6 (18.0-19.0), is arched above the orbits, rises posteriorly, greatest length of skull 14.8 (14.7-15.0), and is said to rise abruptly. Of diagnostic im- breadth of braincase 7.0 (6.0-7. 1). Miller and portance, the arched cranium extends poste- Allen (1928) list skull length and breath of riorly with even a slight upward curvature at braincase as 14.6-15.5 and 6.8-7.6, respec- the lambdoid crest. In consequence, the up- tively. Weights are about 41-43 percent low- per portion of the supraoccipital bulges pos- er in winter. See Table Chi-6. teriorly. There is no median sagittal crest. Dental Formula. 1 2/3 C 1/1, P 3/3, M Myotis lucifugus shows less curvature in the 3/3 = 38 teeth. rise of the braincase, and less posterior pro- Geographic Range. Expected throughout truberance of the supraoccipital. Myotis so- the state and the Upper Peninsula. See Map. dalis shows a comparable rise, but much less Status. Keen’s Myotis is often considered posterior protruberance of the supraoccipi- rare (e.g., Coggins et al., 1981) in Wisconsin tal. See accounts of other Wisconsin bats to (Jackson, 1961, only documented 26 speci- distinguish them from M. keenii. mens). This species is actually fairly common during migration in central Wisconsin, and the numbers examined suggest the status throughout the state is improving. The species is decidedly beneficial because it is insectivorous and helps control insect numbers. It can harbor rabies virus, but has weak jaws and probably cannot bite dogs or people (Baker 1983). It is seldom sent to the Wisconsin State Hy- giene Laboratory as a rabies suspect. Habitats. Myotis keenii ranges throughout the state in woodlands, usually near lake shores and streams. The species often hangs up in trees, tree cavities, or under loose bark, in the summer. In winter it is often found hanging in caves. During migration this species is likely to show up almost anywhere, even hanging onto brick buildings, and entering them on occasion. Maternity colonies are usually Skull of Myotis keenii. small, containing up to 30 pregnant females,

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 115 and are found in barns, behind wooden shin- gles on old houses, or under loose tree bark. Whereabouts of the males are open forest lands above shrubby understory, or over streams. In winter, Keen’s bats are occasionally found in caves with other cave-dwelling bats, especially in southwest Wisconsin. They seldom exceed 10 animals in a hibernaculum, but might number as many as 300. They prefer cool, moist and quiet areas in a hibernaculum. Hibernation may last for eight to nine months (Fitch and Shump 1979). Foods. Insects and other small invertebrates are eaten. LaVal and LaVal (1980)

Maps showing geographic distribution of Myotis keenii in Wisconsin and also in North America.

116 THE WILD MAMMALS OF WISCONSIN found Keen’s bats to prefer small moths, cad- Additional Natural History. Fitch and disflies, stoneflies, homopterans, mayflies, Shump (1979) reviewed biology for M. kee- Foods. Insects and other small invertebrates nii. are eaten. LaVal and LaVal (1980) found Specimens examined. Total, 51. Bay- Keen’s bats to prefer small moths, caddisflies, field, Brown, Crawford, Dodge, Grant, Iowa, stoneflies, homopterans, mayflies, beetles, Iron, Jackson, Lafayette, Monroe, Pierce, and flies. Rarely mosquitoes are ingested. Portage, Richland, Washburn counties. Reproduction. There seems to be winter Michigan: Big Summer Island, in Green sperm storage in Myotis keenii. One young, Bay 1. carried in late May or June, is apparently born in June and rapidly develops (Hoffmeister 1989, Kurta 1980, Clair et al. 1979, Easter- Myotis lucifugus lucifugus LeConte la, 1968) until weaning about 32 days later Little Brown Myotis or Little Brown Bat (Kunz 1971). Lactation occurs usually from June to August, but usually not later than July. 1831. V [espertilio]. lucifugus LeConte. In McMurt- A juvenile from Sparta, Wisconsin (UWSP- rie, The Animal Kingdom... 1:431, type from 7878) was taken 11 May 1995, and an em- Georgia. bryo (27 mm) was found 24 June 1980. 1897. Myotis lucifugus: Miller. N. Amer. Fauna, Keen’s bats are usually considered adults 13:59. when they show fusion of the epiphyses of the wing, a dubious way to predict sexual The generic name means “mouse- maturity (Long and Jones 1966). This meth- eared.” The word lucifugus means to hide od does appear to segregate young which from the light. have not attained full growth from old adults Description. The little brown bat is small, that live up to 18 years (Barbour and Davis with ears neither broad nor elongate, with 1969). tragus neither pointed and high nor blunt and Mortality. No information is available on short. It lacks a keel on the calcar except rare- fleas, bugs, ticks, and chiggers parasitize them ly, and its elastic fibers of the uropatagium (in South Dakota). Intestinal and other worms are numerous, crowded together, and usually are also reported, by Whitaker and Mumford more than 18 lines (see fig. and account of (1971) and by Coggins et al. (1981) for Wis- Myotis keenii). The skull is not flattened, but consin. the rise of the cranium posteriorly is gradual, Home Range and Density. Little infor- and the occiput is neither high nor does it mation is available. Hoffmeister (1989) reports bulge much posteriorly. There is no sagittal that in winter 400 to 800 estimated individu- ridge, and the palate is broad across the mo- als were congregated in the Black Ball Mine, lars. Hairs regularly extend beyond the claws Illinois. No densities even approaching this of the hind toes. number have been seen in Wisconsin. Beer The number of chromosomes is 2N = and Greeley (in 1947) banded more than 100 44, FN is 50 as in all Myotis (Fenton and in an iron mine near Hurley, Wisconsin. Barclay 1980). Geographic variation. There is no geo- The little brown bat is dark or slightly graphic variation in this species in Wisconsin yellowish brown, the tips of the hairs with a and Upper Michigan. The nominate race, in metallic, brassy sheen, occasionally duller as the Pacific Northwest, is darker, the skull in M. sodalis, and occasionally (when shorter, and the ears more recurved (scimi- bleached) almost blond. The venter is gray tar-shaped). overlain with pale grayish or brownish tips of the hairs. The ear pinnae and the patagial

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 117 membranes are slate gray to nearly black. crease from 329 (54.6 percent) in northern Walley (1971) found an albino in southwest counties to 10 (1.7 percent) in southern coun- Wisconsin. ties. Across Lake Michigan on the Lower Females average a little larger than males. Peninsula, Kurta et al. (1989) showed a drop See table Chi-6. from 141 (62 percent) in northern counties Dental Formula. 12/3,C 1/1, P 3/3, M to only 7 (4 percent) in the southern ones. 3/3 = 38. The big brown bat Eptesicus showed a con- Geographic Range. Found throughout verse trend. M. lucifugus may be the most Wisconsin, but usually congregated in suitable abundant bat in Wisconsin. It surely is in the caves and mines in the winter. Common in caves and mine tunnels in winter, when some southern latitudes. Clark et al. (1987) found populations may exceed an estimated 10,000- in four summers of netting bats in Iowa a de- 20,000 bats. Status. Unlike many of the Wisconsin bats, the little brown bat becomes less common southward Nationally, the numbers of Myotis lucifugus reportedly declined (Fenton and Barclay 1980), probably due to the use of pesticides. There is no obvious decrease in Wisconsin. Habitats. Little browns frequent woodlands, often foraging in riparian habitats or near the mouths of caves. In migration the bats turn up almost anywhere, even entering buildings or hanging onto the exteriors. In winter the bats search out caves hibernacula in which to sleep. Fenton and Barclay (1980) list three kinds of roosts: day, night, and hibernation sites. Day roosts are in buildings, tree cavities, rocks, wood piles, and occasionally caves. Non-reproductive females and males are more solitary. After initial foraging, sometimes bats pack together in confined places for night roosting. Man-made tunnels (mine shafts, storm sewers), old houses, cabins and barns are often used as maternity and temporary Skull of Myotis lucifugus. roosts. Hibernacula are usually caves or mines to which the bats migrate in autumn. Hiber- nation may last from early September until mid-May. Torpor patterns were discussed by Kurta (1991). Distances from nursery colonies to hibernacula are as far as several hundred kilometers. Foods. Gould (1959) reports the little brown bat can capture about 500 insects per hour. Whitaker (1972b) found that 16 specimens had eaten mostly moths, caddisflies, A skull variant of M. lucifugus. flies, leafhoppers, and beetle larvae- usually

118 THE WILD MAMMALS OF WISCONSIN small insects less than 10 mm long. Other insects taken include mayflies, wasps, and stone flies (Barbour and Davis 1969, Belwood and Fenton 1976; Freeman 1981, Anthony and Kunz 1977). Fenton and Barclay (1980) report heavy feeding on aquatic insects. Chi- ronomids are a staple food. Effects of pregnancy and lactation on energetics have been discussed by Kurta et al. (1989). Reproduction. In late summer and fall, the little brown bats often congregate in large numbers, a phenomenon called “swarming.” Fen- ton (1969) found two phases of swarming in

Maps showing the geographic distribution of Myotis lucifugus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 119 Ontario and Quebec. Sexual activity began bors the rabies virus. Rabies is not a signifi- following the August phase of swarming, in cant threat to people from this bat. Less than September. Sperm is stored during hiberna- 1 percent of this species harbor rabies, and tion until spring. Copulation may occur in caves they seldom transmit it to carnivores (Trima- when hibernating bats arouse periodically from rchi and Debbie 1977, Trimarchi 1978). their sleep. Ovulation and fertilization take Home Range and Density. Little infor- place about the time bats leave the hibernacu- mation is available for Wisconsin Myotis lu- la. A single fetus is carried. Maternity colonies cifugus. Jackson (1961) mentions that Beer are formed in abandoned farm buildings be- and Greely (in 1947) observed more than 300 hind boards and timbers, in hollow trees, and little browns in a mine tunnel near Hurley. under bark of dead and dying trees. Long and Theiss (1996 unpublished) report- After nearly two months gestation, birth ed many thousands in hibernacula of Upper takes place on the Upper Peninsula of Mich- Michigan and Wisconsin. Although less nu- igan either of the pectoral teats. The eyes and merous in large cities, compared to small and ears open in a few hours. By late July to early medium-sized towns, M. lucifugus was occa- August weaning takes place, and the young sionally observed in big cities. It is rarely en- can fly at this time, about three weeks after countered in the winter, and seems at peak birth. Permanent teeth are almost all erupt- abundance in August. It is, of course, numer- ed. Females may become pregnant in their ous in the hibernacula. first year. Some little brown bats live 24 years Remarks. One specimen (UW-SP # or longer (Griffin and Hitchcock 1965). The 2749, see Fig.) resembles little browns from longest records are 34 and 35 years (Wayne nearby sites except the ear pinnae were more H. Davis, personal communication). Breed- recurved and grayer in color. The skull was ing over many years may compensate for sin- highly unusual in form, not rising gradually in gle offspring each year. a low, flattened profile, but rising abruptly as Adams (1992) studied the osteogenesis in such forms as Myotis californicus or Myo- of the fetus of the little brown bat, finding tis austroriparius. Perhaps as a consequence that the appendicular long bones ossified ear- of the abruptly rising cranium, the skull was liest of the postcranial skeleton, probably be- very short in total length (13.7mm). Such cause bats need their wings for flight. Adults abrupt curvature is seen in other species un- are distinguished to some extent by toothwear, known in Wisconsin. The specimen was sent and young from older adults by fusion of the to Dr. Wayne Davis, an authority on bats, who epiphyses in the bones of the wing in the lat- identified it, unusual features notwithstanding, ter (see Burnett and Kunz 1982). as M. lucifugus. Two more Wisconsin M. Mortality. Snakes, raccoons, and house lucifugus, with similarly short and arched cats feed on little brown bats (Long 1978, skulls were seen in the University of Minne- Barbour and Davis 1969, Fenton 1970). Birds sota (Bell Museum) collection. of prey occasionally take them, as do some Geographic Variation. No significant opportunistic smaller birds such as red-winged geographic variation was noted. A great deal blackbirds and common grackles (Garber of individual variation was observed in little 1977, Fenton and Barclay, 1980). brown bats. Parasites include fleas, chiggers, flukes Specimens examined. Total, 283. Adams, and tapeworms (Timm 1975, Scharf and Ashland, Barron, Bayfield, Burnett, Stewart 198, Font 1978). Insecticides may Chippewa, Columbia, Crawford, Dane, cause significant mortality (Kung et al. 1977, Dodge, Door, Douglas, Eau Claire, Fond du Geluso et al. 1976, Gillete and Kimbroug, Lac, Forest, Grant, Green, Green Lake, Iowa, 1970). Occasionally a little brown bat har- Iron, Jackson, Jefferson, Juneau, La Crosse,

120 THE WILD MAMMALS OF WISCONSIN dalis that might identify it if one could see them. They are invisible in the darkness or poor light of a cave. The pelage is dull with less of a rne- tallic sheen on the fur; the venter is duller also, the fur having a slight purplish suffusion; and some hairs that grow between the toes seldom extend beyond. In the U. S. National Museum collections a few labelled sodalis have hairs extending beyond the toes. Showing annual curves for abundance of M. lucifugus for 1992-1994. State-wide samples from state Hygiene Colonies of Myotis sodalis are conspic- Laboratory area mainly based on people to bat encounters, uous in the southern part of its range. The and not on bats observed in caves, the usual source. bats crowd and huddle together, one bat deep, probably to conserve body heat. Such bats are easy to segregate from other closely re- Lafayette, Langlade, Lincoln , Manitowoc, sembling species, without much examination Marinette, Marathon, Milwaukee, Oneida, of subtle characters. An undergraduate re- Outagamie, Pepin, Pierce, Price, Portage, search student, Melanie Jones, pointed out Richland, Rock, Shawano, Sheboygan, St. to me a new character that distinguishes the Croix, Taylor, Trempealeau, Vernon, Vilas, little brown bats from the Indiana bat, at least Walworth, Washburn, Waukesha, Waupaca, from my few (11) specimens of M. sodalis. Waushara, Wood counties. The hind toes of M. lucifugus are larger, ob- Michigan. Delta Co.: Big Summer Island viously thicker and with longer claws than in 7. Poverty Island 5. St. Martin’s Island 2. M. sodalis. Generally the toe length failed to attain lengths of 5 mm in sodalis, and usually exceeded 5 mm in lucifugus. If this character Myotis sodalis Miller and Allen is good, then perhaps it will be easier to iden- Indiana Myotis or Indiana Bat tify the two species in dark caves. See the aforementioned comments about 1928. Myotis sodalis Miller and Allen. Bull. U.S. duller dorsal pelage and purplish suffusion on Nat. Mus., 144:130. Type from Wyandotte the venter of Myotis sodalis. The hairs, on Cave, Indiana. very close examination, seem somewhat tricolored (not distinctly so as in pipistrelles) in The generic name refers to “mouse- that the basal portion of each hair is black, eared” bats, and the second name suggests the becoming gray in the middle, and the hair is huge communal masses of these bats hanging tipped with a tiny segment of brown. up in some caves. Karl Koopman (personal Myotis sodalis resembles Myotis lucifu- corr.) suggested that sodalis means “associat- gus in size. See Table Chi-6. ed” with M. lucifugus. If he were correct, the Dental Formula. 1 2/3, C 1/1, P 3/3, other usage seems more meaningful. M 3/3 = 38. Description. One reason the species is Geographic Range. The known occur- represented so poorly in mammal collections rence of Myotis sodalis in Wisconsin is based is the failure to recognize the species. It close- upon a single specimen taken in a huge hi- ly resembles other brownish, medium-sized bernaculum called Beetown Mine (or Atkin- bats. Field collectors usually do not kill every son’s Diggings). Repeated attempts to collect animal encountered, but collect selectively, and another specimen from this mine and nearby with ecological considerations. There are three caves have so far proved fruitless. Netting and cryptic characteristics of the living Myotis so- searching mines during the summer in south-

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 121 of its range, because rare species are not usually mentioned unless their residence is well known. In my opinion, the species should be placed on the Wisconsin Endangered Species List, even if it is peripheral. Habitats. The Indiana bat moves into caves in winter (from November to April). In spring, maternity groups of a few females roost in outlier homes (i.e., temporary shelters away from the main home), behind loose bark, under bridges, in hollow trees, and occasionally in buildings. Humphrey et al. (1977) studied the summer habitats of Myotis sodalis. This species may form nursing colonies in green ash trees (Kurta et al., 1993). Another nursing colony was found in a Michigan sycamore (Kur- ta et al., 1994). They also occur in oaks (Quer- cus), elm (Ulmus), and hickories (Carya) (Th- ompson, 1982). This species often forages over slow-moving creeks and streams lined with Skull of Myotis sodalis. trees (Kurta, 1980). Clark et al. (1987) found them in agricultural fields and woodlands (dis- west Wisconsin for two summers also yielded turbed riparian woods, forested floodplains and no specimens (Ainslie 1983). Nevertheless, upland forests) in Iowa. this species is known in several counties in There are three kinds of roosts. Hibe- southern Michigan, in Cook County, Illinois macula caves are usually found south of Wis- (near southeast Wisconsin), and from Joe consin. Nursery roosts and outliers are the Davies County, Illinois (4 specimens), just other homes. There are also temporary out- south of Beetown Cave. The species is rare liers in tree hollows and under tree bark (Kur- but present in northeastern Iowa, although ta et al. 1 993). Maternity colonies, up to 28 no reproductive colonies have been found. females but no males, are often found under The species is an uncommon visitant in Wis- bark of trees. consin but also a possible resident in south- Foods. Moths seem preferred over small ern Wisconsin from time to time. So little is beetles, wasps, bugs, flies, and caddisflies (La- known of the maternity roosts (Kurta et al., Val and LaVal 1980; Whitaker 1972). About 1993, 1994) that in Wisconsin they may have 90 percent of the food of Myotis sodalis is been overlooked. comprised of soft-bodied insects, and there is Status. The Indiana Myotis is listed as a a preference for moths by lactating females. federal Endangered Species. In some places In fall, diverse insects are eaten. the numbers of those bats declined as much Reproduction. In late summer in the as 28 percent in 15 years (Thomson, 1982). breeding range of Myotis sodalis, breed- Because this species hibernates in vulnerable ing swarms assemble in the evening sky, and masses that are frequently vandalized by hu- mating occasionally continues in hibernac- mans visiting caves, this species is also includ- ula in September and October (Cope and ed also on the Illinois State Endangered Spe- Humphrey 1977, Mumford 1969, Thomp- cies List. It is not included on the Wisconsin son 1982). The female bats hibernate, with list, although rarer in Wisconsin than in most fertilization delayed until emergence from

122 THE WILD MAMMALS OF WISCONSIN the hibernacula in April (Hall 1962, LaVal and LaVal 1980). As in other vespertilion- ids, the female contains a gelatinous plug holding the sperm until it dissolves in spring. Ovulation and fertilization seemingly occur at this time. A single fetus is carried by each mother, from May 20 to June 24 (Hall 1962, Easterla and Watkins 1969). Lacta- tion follows parturition from late June to mid-August (Kurta 1980), when the young begin flying about with the adults. Clark et al. (1987) found pregnant females as late as late June in southern Iowa.

Maps showing geographic distribution of Myotis sodalis in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 123 Clark et al. (1987) found no evidence of Genus Lasionycteris Peters breeding in northern Iowa, although breeding was common in southern counties. They Lasionycteris noctivagans (Le Conte) suggest the flight distance from hibernacula Silver-haired Bat in central Missouri taxes pregnant females moving northward (about 463 km more or 1831. V [espertilio]. noctivagans Le Conte. In Mc- less) to establish nursery roosts (the reproduc- Murtrie. the Animals Kingdom... Baron von tive females taken were netted). Bowles (1975) Cuvier. 1:431. Type from eastern United States. had taken hibernating males and torpid fe- 1865. Lasionycteris noctivagans: Peters Monatsb. males in northeastern Iowa, not far from the Preuss Akad. Wiss. Berlin, p.648. hibernating Wisconsin specimen. Time might be a factor, for a pregnant female approach- Lasionycteris noctivagans noctivagans ing parturition probably cannot postpone it, (Le Conte) and after leaving the hibernacula females must seek out opportune nursery sites. Kurta et al. The name Lasionycteris means hairy bat, (1993) found reproductive females and young and noctivagans means night wanderer. There (45 bats) together on green ash trees (Fraxi- is no silver on this bat, but the cream or white nus pennsylvanica), in southern Michigan. wash on such dark fur appears silvery white. Indiana Bats are aged primarily by clo- Description. This medium-sized bat has sure of the epiphyses of the bones of the wing. fur extending from the back onto the basal Old bats may be recognized by toothwear. One part of the uropatiagium, occasionally more banded bat lived 20 years (Thomson 1982). than half way to the border. The tragus is Mortality. Few predators are known ex- exceptionally short (5-9 mm) and blunt, and cept black snakes Elaphe obsoleta (Barr and the calcar is not keeled. The tail is short (less Norton 1965), mink and screech owls. Mites, than 40 percent of total length). The skull is flukes and tapeworms parasitize this species flattened (see Fig.). The baculum is large, and (Whitaker and Mumford 1970; Thomson, the karyotype is distinctive in the family, 2N 1982). No rabies virus has been reported in = 20, FN = 28 (Kunz, 1982). M. sodalis. People may kill them when they The fur is basically blackish brown, over- find the communal clusters in caves. lain with white- or cream-tipped hairs. This Home Range and Density. The clusters “frosted” effect is why the bat is called “sil- on cave walls, in the coolest parts of the cave, ver-haired.” The ears are black. Older bats are huge but only one tier deep (Thomson may lack the whitish frosted tips of the hairs, 1982). No information for clusters is avail- and some are even yellowish (Kunz, 1982). able in Wisconsin. Humphrey et al. (1977) Measurements are given in Table Chi-6. found home range to be small. Maternity sites Weights vary up to 11 g (Kunz, 1982). certainly may be expected in Wisconsin, es- Dental Formula. The silver-haired is the pecially if the species ever builds up its num- only Wisconsin bat with 36 teeth. It has only bers. In Iowa it is only 5.6 percent of the total two pairs of upper premolars; there are three bats netted in summer, and is, as in Wiscon- lower premolars on either side. DF = I 2/3, sin, quite rare in winter. C1/1, P 2/3, M 3/3 = 36. Additional Natural History. Thomson Geographic Range. To be expected in any (1982) reviewed biology of Myotis sodalis. county throughout the state. This bat seldom Geographic Variation. There is no evi- winters in Wisconsin (one record, Schmidt, dent geographic variation. 1931). Furthermore, it has seldom been taken Specimen examined. Total 1. Grant in any parts of Wisconsin except along the for- Co.: Beetown Cave, 1 UI. ested Lake Michigan western shore. Apparent-

124 THE WILD MAMMALS OF WISCONSIN ly in migration they congregate along the shore Habitats. The silver-haired bat prefers and wend their way southward. Elsewhere in riparian areas along fast-moving brooks or Wisconsin, the bat and its habits are poorly streams in summer, and often is found hang- known, and it seems one of the rarest of our ing up to rest in woodpiles, hiding behind the mammals. Approximately 10 percent of bats loose bark of tree or in tree cavities. Schmidt netted four summers in Iowa were of this spe- (1931) found one on 6 September, in Clark cies (Clark et al. 1987). In a state-wide sample County, in his basement under the furnace. of bats sent from the Wisconsin Hygiene Labo- Long and Thiess (unpublished) record some ratory, collected from 1991-1995, this species specimens for Wisconsin in October. Frank was also found in northern and western Wis- Iwen and Scott Craven captured one in Mad- consin (Long and Theiss, unpublished) (see ison on 15 Dec. 1981. Silver-haired bats hi- Map). Dates vary from April 21 to October 19 bernate in caves and mines in southern Illi- in this collection. In Iowa it is only 5.6 percent nois and other states south of Wisconsin. of the total bats netted in summer, and, as in Foods. The diet is comprised chiefly of Wisconsin, is quite rare in winter. moths, flies (Freeman, 1981; Kunz, 1982) and Status. See Range. This rare and beauti- emerging aquatic insects. This bat is known ful, silvery bat is now considered dangerous as a “moth strategist,” but feeds on several as a rabies threat throughout the United diverse insects. The energy budget per day States. Rabies viruses can be identified by was on average 4.86 Kcal. antibodies in the victim, and the silver-haired Reproduction. Testes enlarge in late July bat has been implicated even when it was through November (Turner 1974). Very like- unseen, even if its biting was never observed. ly sperm storage occurs in females during their This implication certainly needs further inves- hibernation. Kunz (1982) lists a brief gesta- tigation. Rabies has diminished in some oth- tion of only ten days. Ovulation peaks in April er animals, e.g. dogs and cattle. and early May, with implantation delayed about 50 days. Females with one or two embryos were collected in late May and June. One silver-haired bat captured alive gave birth to two offspring on June 20 (Kurta and Stew- art 1990). During this birth the female sat upright, the uropatagium curled up like a bas- ket. The first young was born breech, lacking its amnion. The young, a female, pushed against the mother with its hind legs, while the mother constantly licked her. Birth was complete in 23 minutes, and the mother continued to lick the chirping infant. The newborn found its way to the mother’s left nipple. The placenta had not been passed. The second newborn began to emerge breech, about five minutes later. Again, the amnion was already ruptured. This young (a male) did not push against the mother, but it too was licked. In 16 minutes this birth was complete. The mother dragged the chirping male, and this pulled forth its placenta. The other pla- Skull of Lasionycteris noctivagans. centa then was extruded. The mother ate them

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 125 both; she did not sever the umbilical cords. The young male’s wing was entangled by the cord, and it did not reach a nipple. The cord was stretched but still red in appearance 33 minutes after birth. The mother weighed 11.0 g, and the young 2.5 g each. The baby female was 44.7 mm total length, the male 47.1. Kunz (1982) records weights of naked, newborn young as 1.9-1.8 g. The young are pink, with mostly tan and black wings, the skin naked, the eyes closed. In three weeks the young are capable of flight and of catching insects for themselves. One Wisconsin female (UW-SP 7161) taken on April 21,

Maps showing distributions of Lasionycteris noctivagans in Wisconsin and North America.

126 THE WILD MAMMALS OF WISCONSIN contained two embryos measuring 7 mm in Perimyotis subflavus subflavus (F. Cuvier) crown-rump length. Some bats live 12 years. Mortality. Little information is available Baker (1983) translated Pipistrellus as “bat”, on the silver-haired bat’s enemies. Sperry and because of its diminutive ending I sup- (1933) reported striped skunks (Mephitis) pose the word meant “little bat.” Since the preying on them during their hibernation. little bats are no longer regarded as true pip- Owls eat them (Kunz, 1982). Parasites are istrelles, neither in the East or West, I resur- listed by Kunz (1982) and include mites, bat rect Georgia, for the type locality and former flies, a flea, two species of Cimicid bat bugs, usage. The word subflavus means a little less a nematode, seven trematodes, and three than yellow, meaning either pale yellow or cestodes. Rabies is often reported from this yellowish. bat, especially in young of the year. In the Description. This tiny pipistrelle bat with beginning of these accounts of Chiroptera are short ears and blunt tragus has a long tail remarks on the importance and prevalence (about half the total length). The calcar lacks of the strain of virus associated with this mi- a keel. The skull is tiny, delicate, with broad gratory, woodland bat. rostrum and inflated braincase. The braincase Home Range and Density. Seldom are is neither flattened nor particularly high in these solitary bats (Kunz 1971, 1982) caught lateral profile. There are sparse hairs along in groups larger than two or three, even in the basal margin of the uropatagium. hibernation. This blond bat with conspicuous black Additional Natural History. Kunz (1982) wings (showing red blood vessels and pink reviewed natural history of Lasionycteris bones in life) is exceptionally beautiful. The noctivagans. dorsal hairs, on close examination, are dis- Geographic Variation. No significant geo- tinctly tricolored, dark gray basally, buffy sub- graphic variation has been noted in this species. terminally, then brown or reddish brown. (To Specimens examined. Total, 24. Adams, see this trait, blow on the fur and fan it out in Bayfield, Brown, Dane, Door, Kenosha, a circular pattern, bare in the center.) The Manitowoc, Milwaukee, Outagamie, Ver- venter or belly is also yellowish brown. The non, Waukesha counties. forearm is reddish blond, and the patagial membranes and ears of dried specimens are dusky brown. Genus Perimyotis Menu This tiny bat is significantly smaller even Pipistrelle-like Bats than Myotis lucifugus. See Table Chi-6. Dental Formula. 1 2/3, C 1/1, P 2/2, Perimyotis subflavus (F. Cuvier) M 3/3 = 34. Georgian Pipistrelle Geographic Range. Found normally in southwestern Wisconsin, in the caves of the 1832. V [espertilio]. subflavus F. Cuvier. Nouv. Ann. numerous bluffs there, the Georgian pipist- Mus. Hist. Nat. Paris, 1: 17. Type locality Le relle has wandered north as far as Iron Coun- Conte Plantation, Liberty Co., Georgia. ty and eastward to southeast Wisconsin. Even 1897. Pipistrellus subflavus: Miller. N. Amer. Fau- in summer the species seems restricted to the na, 13:90, published as P. s. obscurus. See hilly country of southwest Wisconsin, and also Hall and Dalquest, 1950:600. migrates to the caves, mines, and sink holes 1959. Pipistrellus subflavus subflavus: Davis. J. for hibernation. Mamm., 40:521-531. Status. This tiny pipistrelle is uncommon 1984. Perimyotis subflavus: Menu. Mammalia, 48: in southwestern Wisconsin, and is exception- 410. See Horacek and Hanak, 1985-6. ally rare in most of Wisconsin. Furthermore,

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 127 productive females in leafy canopies of deciduous trees, even in groups or colonies. Of 37 roost trees, the pregnant or lactating bats were usually hiding in dead foliage. Since upland oak trees were preferred, the pipistrelles were found often on high ground. Foods. The diet consists chiefly of leafhoppers and tiny insects that inhabit the leafy outer branches of deciduous trees. Moths, bugs, tiny beetles, small wasps, and flying ants are preferred foods (Whitaker 1972; LaVal and LaVal 1980). Reproduction. Males outnumber females in caves, a pattern seen also in other hibernating bats. Copulation seems to precede hibernation; the sperm is stored in the female, and is viable until spring. Mating may also take place in spring according to Baker (1983). Two young are born from mid-June to early July. Usually the young are sheltered Skull of Perimyotis subflavus. in dense tree canopies, bark crevices, tree cavities and probably rock crevices. After ap- its numbers seem to be decreasing in the hi- proximately 30 days the young are weaned bernacula. This bat should be protected im- and can fend for themselves. Pipistrelles live mediately especially in caves, possibly secured at least 10 years (Baker 1983, Davis and during hibernation by iron gates. Caves should Mumford 1962, Davis 1966). be set aside to protect it and the peculiar cave Mortality. There is little information on the faunas associated with bats. The bat is bene- Georgian pipistrelle’s enemies. A leopard frog ficial because of its insectivorous habits, and ate one (Creel 1963). Owls and other birds feed it is esthetically attractive owing to its tiny size, on summer bats, whereas snakes, raccoons, blond fur, and beautiful black wings. No spec- skunks and other predators occasionally cap- imen up to now has been found to harbor ture adults. People often kill them during hiber- rabies virus in Wisconsin, although some have nation, and even field collectors may decimate in other states. But these bats probably can- the low populations in Wisconsin. Whitaker and not bite through the skin of humans, dogs, or Wilson (1971, 1974) report mites, chiggers, and raccoons, because the jaws are minute and flukes parasitizing pipistrelles. weak. The bat might be regarded as a vector Home Range and Density. In summer, or source of the virus. pipistrelles are seldom seen, but may form Habitat. The Georgian pipistrelle is found small groups. Jackson (1961) believed they in deciduous woods generally at the forest roosted in shallow rock crevices in summer. edge, along streams and rocky outcroppings, No information on Wisconsin summer roosts mostly in the hills of southwest Wisconsin. It is available. In southwest Wisconsin Ainslie hibernates in mines and caves. In the Stevens (1983) netted only seven specimens one sum- Point collection are numerous specimens from mer. In winter hibernacula in Wisconsin and caves, but very few specimens have been tak- Upper Michigan, pipistrelles are never numer- en in summer. J. I. Veilleux (2001) reported ous (Long, personal observations; Stones and that in Vigo County, Indiana, he found re- Haber 1965; Greeley and Beer 1949).

128 THE WILD MAMMALS OF WISCONSIN Additional Natural History. Fujita and Kunz (1984) reviewed biology for this species, as Eastern pipistrelle. Geographic Variation. The four races in P. subflavus vary slightly in size and color.. Wisconsin pipistrelles belong to the nominate race (Davis, 1959), and show no geographic variation. Specimens examined. Total, 73. Craw- ford, Dodge, Grant, Iowa, Lafayette, Pierce, Richland, Sauk, Vernon counties. Other Records. Upper Mich. Baker (1983): Keweenaw and Ontonagon counties.

Maps showing geographic distribution of Perimyotis subflavus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 129 Genus Nycticeius Rafinesque The evening bat is brown, with ashy-brown sheen on the dorsal fur, but resembles M. lu- Nycticeius humeralis (Rafinesque) cifugus, M. keeni, and E. fuscus. It has less Evening Bat brassy gleam of the dorsal fur. The grayish venter is tinged creamy buff. The wings and ears 1818. Vespertilio humeralis Rafinesque. Amer. are black. Measurements are in Table Chi-6. Monthly Mag. 3:446. Type from Kentucky. Dental Formula. Unique among bats in 1819. N [ycticeius]. humeralis: Rafinesque. J. Phys. Wisconsin. 1 1/3, C 1/1, P 1/2, M 3/3 = 30. Chim., Hist. nat. et Arts, Paris, 88:417. Geographic Range. Ordinarily one would 1883. Scotophilus carolinensis: Strong. Geology of not list a species from Wisconsin without a spec- Wisconsin, Survey 1873-1879. P. 436-440. imen. Almost all range maps I have seen published for the species shade in southern Wis- [Included with brown, silvery, and Geor- consin as part of the probable range. There is gia Perimyotis (=Pipistrellus) subflavus) bats, one record (see below) from 1883. The species all four were given separate names under Sc- is known (Hoffmeister, 1989) from three sepa- otophilus. The name Vespertilio carolinen- rate northern Illinois localities (Winnebago, Lake, sis E. Geoffroy-St. Hilaire was to be placed and northern Cook counties) bordering Wiscon- in Eptesicus, not anticipated by Strong, of sin. The northern wanderings must extend a course, and possibly not even synomynized little northward of these peripheral localities. The correctly by Elliot (1907). Hollister (1910) mean of three homing distances for Nycticeius suggested that Strong’s carolinensis was given by Watkins is 96 km (60 miles) and the Nycticeius, often known as Scotophilus, lat- longest distance homed was 153 km (94 miles). er documented from several Illinois specimens Lack of Wisconsin specimens may be due to taken near Racine. See comments below.] the difficulty in distinguishing this bat from little brown bats. In summer, little collecting has been

Nycticeius humeralis humeralis (Rafinesque)

Nycticeius means nocturnal. The name humeralis means arms or wings. Watkins (1972) considers the meaning as nocturnal, and per- taining to the humerus. Description. The species closely resembles Little brown and small Big brown bats (which are usually larger). Externally the muzzle of Nycticeius is dark, and the ear tragus is very short and rounded (arcuate). The calcar is not keeled. The skull of the evening bat differs markedly from either of the aforementioned, being shorter, broader, and flattened. There are only 30 teeth, and the single upper incisor on each premaxillary extension is distinctive. It is closely sited next to the canine. Most vespertilionid bats have two upper incisors on each premaxillary bone. The karyotype is 2N = 24, FN = 48 (Watkins 1972), and surprisingly resembles that of the western pallid bat, Antrozous pallidus. Skull of Nycticeius humeralis.

130 THE WILD MAMMALS OF WISCONSIN accomplished along the southeast border of Mortality. Watkins and Shump (1981) Wisconsin. The Nycticeius do not use caves. suggest raccoons, domestic cats, and black rat Not mapped for state. snakes may prey on evening bats, if any fall to Status. See Geographic Range. the ground. There is little known on enemies. Habitats. Nycticeius prefers trees, even Parasites are mites, bat bugs, roundworms, and lone trees, of green ash and silver maple (Acer tapeworms (Watkins and Wilson, 1974). saccharhinum) (Kurta 1980). Evening bats find Home Range and Density. No informa- places to hide in human habitations, hollow tion is available. In Missouri, nursery colonies deciduous trees, and loose bark. S. L. Veilleux harbor 1,000 or fewer, usually much fewer (2001) reported reproductive females in Vigo individuals (Watkins, 1972). County, Indiana, in the green ash and silver Additional Natural History. Watkins (1972) maple, always below the surrounding tree can- reviewed the biology for the evening bat. opy at heights averaging approximately 13 m. Documented Record. Strong (1883: Such habitat is more or less continuous from 438) reported this bat from extreme south- the Chicago-Rockford urban areas into southeast Wisconsin under the binomen Scotophi- east Wisconsin, and even far north and west. lus carolinensis, which odd name obscured Foods. Nycticeius feeds on aerial insects. the record until I consulted the older litera- Reproduction. The sexes congregate in ture. See synonymy above. autumn, and two (1-2) young are born from mid-May (in the southern states) to mid-June or even early July. Newborn young develop Genus Eptesicus Rafinesque rapidly, moving to the nipples shortly after birth, the eyes opening in one or two days and Eptesicus fuscus (Palisot de Beauvois) hair appearing in five. After 20 days the young Big Brown Bat reportedly can fly (Jones 1967, Watkins 1972). Wings attain adult length in 30 days. Weaning 1796. Vespertilio fuscus Palisot de Beauvois. Cat. takes place in six weeks. Growth and develop- Raisonne du mus. de Mr. C.W. Peale, p 18. ment were described by C. Jones (1967). Type from Philadelphia. 1900. Eptesicus fuscus: Mehely. Monographia chi- ropterorum Hungariae, p. 206, 338. 1969. Eptesicus fuscus fuscus: Long and Severson. J. Mamm., 50:621-624.

Eptesicus fuscus fuscus (Palisot de Beauvois)

Appropriately the name Eptesicus means “fly- er in the house. “The name fuscus means dusky. Description. The big brown bat is large with a broad face and rhinarium (nosepad), long forearm, skull elongate and flattened, showing little rise in the curvature of the cranium, and in dorsal outline skull wedge-shaped anteriorly much as in Myotis (which are smaller). There are two pectoral mammae. Map of North American distribution of Nycticeius Upper parts of the big brown bat are dark humeralis. or yellowish brown, the color of coffee, with

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 131 a metallic sheen. Belly coffee-brown or gray- mammal. Owing to utilization of houses, build- ish brown, occasionally whitish in pale bats; ings, even church basements, this species has ears and patagia slate or brownish gray. A doubtless increased its numbers (but not its pop- few albinos are known (Jackson 1961, and ularity). Baker (1983) suggests they have in- others). The diploid number of chromosomes creased “many fold,” and they are on occa- is 50, PN is 48. Some Wisconsin specimens sion truly pests in human houses. Long and are bleached (Long and Severson, 1969). Thiess (1996 unpublished) observed a dramatic Measurements are given in Table Chi-6. rise of Wisconsin Eptesicus. They are most The big brown bat is slightly larger than ei- common in towns of 40,000 to 100,000 peo- ther the red or little brown bats; large individ- ple, where they are by far the predominant uals are exceeded in size only by hoary bats. species (Table Chi-3, 4). Curiously, they are Dental Formula. 1 2/3, C 1/1, P 1/2, less common in the largest city, Milwaukee. M 3/3 = 32. They seem to have replaced Lasiurus borea- Geographic Range. The big brown bat lis, which was formerly, in early literature, the ranges widely throughout the state, and of- most common reported bat of towns and set- ten may be found hibernating in mines and tlements. Whitaker (1995) requested that states caves, houses, churches and other buildings. and perhaps federal agencies provide protec- The species is more common in the southern tion of these bats and their maternity colonies, and central counties. Most of these, except because they feed on insect pests (see Foods). Iron County, have numerous southern plant Habitats. The typical habitats are decidu- communities and town-farmland habitats. The ous woodlands, as well as urban areas. Big same trend is seen in Lower Michigan (Kurta brown bats seek caves and dark buildings in et al., 1989), where the bat is more common man-made structures and habitations. I have in agricultural and urban habitats. seen them winter, tree cavities and hollows and Status. Aside from harboring the rabies buildings in summer, and during the year uti- virus in a small proportion of big brown bats, lize a variety of in summer hanging onto build- this species is a beneficial insect-controlling ings even hiding in crevices of my chimney. Foods. Big Brown Bats select larger prey items than Myotis, especially beetles (Whitak- er 1972b, 1955; G. Phillips, 1966, Kunz and Fenton 1973, Hamilton 1971). They also feed on bugs, ants, leafhoppers, moths, crane flies, midges, and other insects (Whitaker 1972, 1995), including several serious beetle pests, and stink bugs. Dipterans, Lepidopterans and other kinds of insects were minor foods. More information on diet is given in Coutts et al. (1973). Foraging begins after sundown, and is usually within 2km of the roost. Big brown bats seldom feed November to March even though they are often seen flying about (Hoffmeister 1989). During lactation the females show a higher metabolic rate and produce more carbon dioxide (Kurta et al. 1990). Data from Juvenile Big Brown Bats from Wisconsin are as follows, including Mus. No., Skull of Eptesicus fuscus. Date of Collection, and Locality:

132 THE WILD MAMMALS OF WISCONSIN 7195 July 30 Athens, Marathon Co. 7188 Aug. 3 ElkhartLake,Sheboygan Co. 7218 Aug. 17 Oregon, Dane Co. 7095 June 30 Ellsworth, Pierce Co. 7176 July 13 Big Bend, Waukesha Co. 7177 July 13 Eden, Fond du Lac Co. 7183 July 27 Cadott, Chippewa Co. Reproduction. Yearling males and some yearling females may breed. Copulation takes place just before and after entering hibernacula, and continues during arousals in the winter (Beer 1955). Sperm is stored until April, and gestation lasts about 60 days, until late May to mid-June, when one or two, and rarely three,

Maps of distribution of Eptesicus fuscus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 133 are born. Apparently there is some kind of more numerous (Barbour and Davis, 1969). In uterine control resulting in a single young (Bir- Wisconsin at Twin Bluffs, we observed several ney and Baird 1985). Newborn young are na- hundred big brown bats. Usually only a few are ked, inactive, blind and weigh only about 2-3 g found, co-existing with but outnumbered by oth- (Barbour and Davis 1969, Davis et al. 1968, er bats, especially Myotis lucifugus. Homing Kurta and Baker 1990). Maternity colonies may cover 400 km, taking 4 to 6 days. seem surprisingly rare, but apparently a moth- Long and Thiess (1996 unpublished) er leaves her young hanging and searches for found the numbers to peak in August and them after foraging (Brenner, 1968; Kunz, September, and probably they do not dimin- 1976; Barbour and Davis, 1969). Lactation ish much in the caves. They are collected in continues as late as early August. all months of the year, and probably are more Some details on growth and aging are active than other Wisconsin bats in winter. given by Burnett and Kunz (1982). Big Brown Beer (1955) studied banding records Bats may live 19 years (Goehring 1974, Bar- (1940-1953) for 3,871 big brown bats in Wis- bour and Davis 1969). Specimens are aged consin and Minnesota. They did not make long by toothwear at the tip of the canine. Closure movements (33-61 miles were the longest), of epiphyseal sutures of the wing distinguish- usually less than 10 miles (= 16 km). The mor- es immature from older bats. Christian (1956) tality rate was approximately 40 percent a year. noted steady growth from birth until mid-sum- Additional Natural History. Goehring mer, after which the growth rate diminished. (1972) made a thorough study of Eptesicus By the seventh summer, canines may be worn fuscus in Minnesota. Another important ref- down to the occlusal rim of the cingulum. erence is Kurta and R. H. Baker (1990). Mortality. Owls, American kestrels, com- Geographic Variation. Long and Sever- mon grackles, opossums, long-tailed weasels, son (1969) found only a single race in Wis- striped skunks and snakes feed on big brown consin. Many bats along the western border bats (Beer 1953, Black 1976, Dexter 1978, were exceptionally pale and had creamy whit- C.F. Long 1971, and Long 1978). Trema- ish underparts. todes were reported by Lotz and Font (1983). Specimens examined. Total, 368. Ash- In summer 1995, a bed bug was found on a land, Barren , Brown, Buffalo, Burnett, Cal- dead Eptesicus from Wisconsin. Other para- umet, Chippewa, Clark, Columbia, Crawford, sites include Basilia and Cimex. Acarine par- Dane, Dodge, Door, Douglas, Dunn, Eau asites include Acanthopthirius and others. Claire, Grant, Green, Green Lake, Iowa, Iron, Several species of mites also parasitize Epte- Jackson, Jefferson, Juneau, Kenosha, La sicus. Kurta and Baker (1990) summarize Crosse, Lafayette, Langlade, Manitowoc, hosts of internal parasites. Over 20 helminths infest Eptesicus in Wisconsin alone (Lotz and Fout 1985). Rabies is a problem in Eptesius nationwide. Encephalitus is possibly transmitted. Histoplasmosis may be contracted by this bat or passed in its guano (Bartlet et al. 1982, Hoff and Bigles 1981). Man-made chemicals cause some mortality (Kurta and Baker 1990). Home Range and Density. In spring, female big brown bats congregate in maternity colonies. These are in habitations of people or Showing seasonal abundance of big brown bats based in oak, beech and other trees. The colony is on Wisconsin Hygiene Laboratory specimens. Long and usually 25-75 adults. In hibernacula males are Theiss.

134 THE WILD MAMMALS OF WISCONSIN The name Lasiurus means hairy tailed, and borealis means northern. The name is a mis- nomer because the bat probably has extended its range northward from southern latitudes, but not very successfully into boreal habitats. Description. This medium-sized, pug- nosed bat is easily identified by its reddish furry uropatagium (a portable “sleeping bag”). The skull has the offset premolar in the “crab-like” Sketch of big brown bat. Charles and Elizabeth skull (see description in Genus Lasiurus above). Schwartz, and Univ. Missouri Press. The size is significantly smaller than in L. cinereus (see Measurements). The ear is round- Marathon, Marinette, Marquette, Milwaukee, ed, nearly circular, and the tragus is short and Monroe, Oconto, Outagamie, Pepin, Pierce, blunt. The rostrum is exceptionally short, and Portage, Price, Racine, Richland, Rock, the canines procumbent. There are two pec- Rusk, Shawano, Taylor, Sheboygan, St. Cro- toral mammae in females. The karyotype is ix, Trempealeau, Vernon, Vilas, Walworth, 2N = 28, FN = 48 (Shump and Shump 1982). Washburn, Washington, Waukesha, Wau- The fur is bright brick red-orange in paca, Waushara, Winnebago, Wood counties. males, and yellowish-orange in the drabber females (there are exceptions to this dichro- matism). The young resemble the females in Genus Lasiurus Gray color (Timm 1989). The dorsum is “frosted” Hairy-tailed Bats or Furry-tailed Bats or washed with whitish or cream-tipped hairs. A creamy spot is conspicuous on the elbow. In this genus there is a single upper incisor, The red bat closely resembles L. cinereus but on each side, and the skull has such a wide is more reddish and much smaller. palatal notch the appearance of the skull is Measurements are given in Table Chi-6. crab-like (perhaps “fanged” as in the heads Shump and Shump (1982) list average mea- of spiders). The dorsal uropatagium is dense- surements for ten males as follows: total length ly furred, and is nearly concolor with the dor- 108.9, tail length 52.7, hind foot 7.9, ear length sum. In the specimens I observed of two Wis- from notch 10.5, tragus 7.5, and forearm 40.6 consin species, Lasiurus has a tiny peg-like mm. Females averaged eight percent larger. first upper premolar, which is set off line, sit- Dental Formula. DF = I 1/3, C 1/1, P ed slightly inside the tooth row. Shump and 2/2, M 3/3 = 32. Shump (1982a, 1982b) wrote thorough re- Geographic Range. Occur throughout views for Lasiurus borealis and L. cinereus. Wisconsin, but red bats are not abundant in northern counties. They migrate southward from Wisconsin for the winter, and seldom Lasiurus borealis (Muller) use caves. Red Bat Status. The red bat seems more common in the southern counties (judging from Jack- Lasiurus borealis borealis (Muller) son’s comments, 1961), and all but one of the recently collected (in the past ten years) 1776. Vespertilio borealis Muller. Des Ritters Carl specimens observed by Long and Thiess von Linne... Suppl. 20. Type from New York. (1996 unpublished) were from southeast and 1897. Lasiurus borealis: Miller. N. Amer. Fauna, south-central Wisconsin. The species had been 13:105. increasing in numbers northward (Long,

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 135 1976) probably because of the opening of the shrubs. Only on rare occasions do red bats forests, the planting of deciduous trees, the enter caves or buildings. Leafy branches of erection of numerous street lights and other ash, elms, sycamore and other deciduous trees lighting also (which attract nocturnal insects are the usual day roosts. in numbers). One specimen was taken in the Foods. Moths and beetles were observed Apostle Islands, in Lake Superior. In recent foods of red bats in Indiana (Whitaker 1972). years the numbers of the red bat taken in Crickets are found in their diet (Jackson Wisconsin have declined, judging by their scar- 1961). Bugs, leafhoppers, wasps and flying city in large samples tested for rabies (Long ants, flies, and moths are also eaten (Free- and Thiess, 1996 unpublished). The red bat man 1981). possibly may be replaced by numbers of Ep- Some data on juveniles from Wisconsin tesicus in human cities and settlements, al- and Minnesota are as follows, including Age, though direct competition for foods and roost Date, Measurements, and Locality: sites seems unlikely. The decline of the red bats may follow tree removal from city streets, Juvenile June 11 58-21-7-8 -tr 2 Chippewa Falls due to diseases and construction projects. In 2 Juveniles June 23 Tiny, umbilicus present, eyes shut, Henney MN early literature in Wisconsin the red bat gen- Juvenile June 30 64-22-9-9 Tasca State erally was reported as common, and Eptesi- 3.2 g 1 Park, MN (Nursing mother) cus was uncommon. 3 Juveniles July 14 14-15g Stoddard, WI Habitats. The red bat prefers the open 4 Juveniles? Aug. 11 31-34 Stillwater, MN deciduous woodlands and riparian habitats along streams, primarily in the southern and Reproduction. Red bats mate in late sum- central parts of Wisconsin. The trees they mer and fall (Steuwer 1948). Sperm is stored forage in, hang up in, and leave their young until spring (Barbour and Davis 1969). In April in while flying about are elms, box elders, and May the migratory females return to Wis- oaks, hackberry, ash, sycamore, and various consin, the males later. The gestation is unknown, but embryos are found as early as late April. Young are born in June or even later. The neonate weighs about 0.5 g. The female may be overloaded with as many as four young. She must carry them in flight, usually 3 (range 1-5). Stains (1965) weighed four young that outweighed the mother (24.4 g, 12.9 g), and yet she mangaged to carry these in flight. Eventually offspring are left hanging in deciduous trees. The young can fly after approximately four weeks and are weaned near this age (Barbour and Davis 1969). A lactating female (UWSP-7877) from Kenos- ha was taken 29 June 1995, and a mother with young was obtained in Madison on July 7. Mortality. Blue Jays are common predators of red bats with reports in four states (Bak- er 1983). Hawks, owls, opossums, and striped skunks eat them (Sperry 1933). Echolocation failures apparently cause Skull of Lasiurus borealis. mortality of several sorts, especially impale-

136 THE WILD MAMMALS OF WISCONSIN ment on barbed wire (Van Gelder 1956, Long 1964). Many red bats are killed by cars. In addition to rabies, red bats are plagued by numerous parasites (Shump and Shump 1982). Mumford and Whitaker (1982:193) reported at least four dead red bats drowned by an April storm on Lake Michigan. These were “likely in migration when they encountered the storm.” Timm (1989) found 50 dead red bats in autumn and only two casualties in spring picked up near a Chicago hotel. Home Range and Density. Mostly solitary, red bats often forage in groups. Nichols and Stones (1971) reported red bats in large

Maps showing distribution of Lasiurus borealis in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 137 numbers moving across Lake Superior. Little premolar is set inside the tooth row. The ear is known of real density, home range, and pinna is rounded, nearly circular, with a short migration patterns. Dates of collection in my blunt tragus. There are 28 chromosomes (fun- specimens vary from April 11 to October 18. damental number of 48) (Shump and Shump, The species is more likely to be observed in 1982). There are two pectoral mammae. autumn (Long, 1976). The fur is brownish, intermixed with Remarks. The red bat harbors rabies vi- blackish and reddish (sometimes yellowish) and rus occasionally, and when infected it presents frosted with whitish or pale buff. There is a a problem because it may come into contact buff spot near the elbow, and the fur of the with children (Whitaker et al. 1969). This spe- uropatagium is nearly concolor with the back cies is most likely to be found sick in one’s (often washed with silvery gray). The individ- yard and, of course, may bite one’s hand if ual hairs of the back typically have four bands, picked up. Warn the children. basally dark, then yellow, then brownish-black, Geographic Variation. One subspecies then whitish distally. The overall effect dor- occurs in the eastern states, and in Wisconsin. sally is brown, red-brown (nearly maroon), or Specimens examined. Total 42. Ashland, yellowish brown fur frosted with white, which Chippewa, Clark, Columbia, Dane, Jeffer- coloration is unique for Wisconsin bats. The son, Kenosha, Manitowoc, Menominee, Mil- ears are brown. Often flying hoary bats may waukee, Portage, Racine, Richland, Rock, be identified by their swift, direct flight. Mea- Sauk, Shawano, Vernon, Walworth, Wash- surements are given in Table Chi-6. ington, Winnebago, and Wood counties. Dental Formula. I 1/3, C 1/1, P 2/2, M Other Record. Door Co.: Washington 3/3 = 32. Island (Long, 1978). Geographic Range. To be expected throughout the state, particularly during migrations. Formerly thought to reside in the north- Lasiurus cinereus (Palisot de Beauvois) ern coniferous forests, actually the hoary bat Hoary Bat

1796. Vespertilio cinereus Palisot de Beauvois. Cat. du mus. de Mr. C.W. Peale, Philadelphia. P. 18. Type from Philadelphia, misspelled as linereus. 1864. Lasiurus cinereus: H. Allen. Smiths. Misc. Coll., 7(165):21.

The scientific names mean hairy-tailed and gray. “Hoary,” as in the phrase “hoar frost,” refers to the whitish tips of the dorsal hairs creating a frosted appearance. Description. The hoary bat is the largest bat in Wisconsin. The wing span occasionally may exceed 15-17 inches. The combination of size, and brownish fur frosted above and extending onto the entire upper uropatagium identifies this species. The forearm usually exceeds 50 mm. The skull is long and broad, “crablike” in form (wide palatal notch). The canine is large and procumbent, a single incisor, and one tiny Skull of Lasiurus cinereus.

138 THE WILD MAMMALS OF WISCONSIN becomes more abundant in the southern counties and on southward. It is never numerous. Status. This solitary species is observed in small groups in summer (they may be family groups). Unlike L. borealis, this larger Lasiu- rus is rare. In winter it probably leaves the state, so far as it can be determined, and in summer is fairly common in the hilly southwestern counties (Ainslie 1983). Dates of collection in this Museum range from 21 May to 19 Novem- ber, indicating the active period in Wisconsin. Habitats. Jackson (1961) and Long (1977) mentioned that hoary bats are more likely than

Maps showing distribution of Lasiurus cinereus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 139 red bats to occur in coniferous forests. This is the pelage has grown. The young are left hang- true if one means the coniferous forests of ing in foliage at night while the mother forages. northern counties, where red bats are seldom The young bats enter the population in late July, found. Hoary bats generally range through and fly well at 33 days of age (Bogan, 1972; much of the conifer forest of the Rockies, but Shump and Shump 1982). Juvenal males report- Ainslie (1983) in southwest Wisconsin netted edly mate their first year (Druecker 1972). The hoary bats in summer in deciduous trees. This interrelation of reproduction, migration and species is very seldom seen in caves. growth is discussed under the Vespertilionidae Foods. A hoary bat once was seen attack- above (Koehler and Barclay, 2000). ing a Georgian pipistrelle presumably to eat it Mortality. Hawks, owls and snakes prob- (Bishop 1947). The primary food source is the ably prey on hoary bats (Church 1967; Wise- community of flying nocturnal insects, such as man 1963). As in the related red bats, nu- moths, flies, beetles, and hymenopterans (Black merous hoary bats become impaled on the 1972, Freeman 1981, Whitaker 1972). barbs of barbed-wire fences (Denys 1972, Reproduction. Copulation in hoary bats has Hibbard 1963, Iwen 1958, Wisely 1978), and not been observed. This probably takes place suffer other accidents in flight. during the autumn migration either prior to move- Home Range and Density. The hoary bat is ment, during movement, or upon arrival in the rare or uncommon. There is little information on winter range. Males do not seem to accompany densities. The sexes keep separate in summer, the females on their northward migration. By coming together only in the mating season. then, the females are, of course, pregnant, likely Remark. The incidence of rabies in the following a winter period of delayed fertilization. hoary bat is fairly high (Whitaker, 1969). In 1994, Parturition occurs in May and June (Mumford only two Lasiurus cinereus were collected at 1969, Bogan 1972) but these observations were the Wisconsin Rabies Lab, State Hygiene Labo- far from Wisconsin. Usually two young are born ratory, and both had rabies. In wild populations, (1-4) weighing about 4.5 g each. They are ini- the incidence is doubtless much lower. tially cradled in the mother’s wings. A mother Geographic Variation. None is evident with two young (one measuring 88 mm in total in North America. length) was taken in Madison on July 7. There is Specimens examined. 22. Dane, Dodge, a fine silvery-gray hair on the newborn skin, ex- Grant, Milwaukee, Portage, Sauk, Vernon, cept on the naked venter, and in three weeks Wood counties.

Monthly numbers of bats counted in southwestern Wisconsin caves, 1981. Ainslie.

140 THE WILD MAMMALS OF WISCONSIN Table Chi-2Chi-2. A survey of the species of bats in Wiscon- Table Chi-4. By the same survey of bats above, relative sin and their numbers collected by the Wisconsin Hygiene abundance and diversity of bats were determined, related to Laboratories, 1991-1994. All specimens are alcoholic speci- log population size (base 10) for Wisconsin. Milwaukee has mens in the UW Museum of Natural History, and represent the most people, the greatest diversity of bats, and the few- a state-wide survey based on people-to-bat encounters, not est of them in relation to population (Table Chi3). Eptesicus collections with mist nets or from caves. Long and Thiess. and Myotis lucifugus are widely distributed, but uncommon in Milwaukee. The diversity index (D) for the entire state Species Number Percent was 0.5. The three lowest ranked sizes were combined to obtain D = 0.583 for 1.7-2.5, which without combining was Eptesicus fuscus 230 64 Myotis lucifugus 107 29 0.464. Frequency is given for the species obtained. Lasionycteris noctivagans 13 4 Log Eptesicus M. Lasio- L. L. M. Lasiurus borealis 7 2 Size lucifugus nycteris borealis cinereus keenii Lasiurus cinereus 2 0.5 & D Myotis keenii 2 0.5 Other species 0 0 <1.7 6 2 2 Totals 361 100 1.8-2.0 2 1 0 2.1-2.5 6 7 1 Table Chi-3. By use of the above survey the number of & 0.583 bats were related to population size of the communities (from roadmaps). Milwaukee had the fewest bats relative to hu- 2.6-3.0 15 9 0 man population (and its great ecological disturbance). Bats & 0.464 were often collected in towns and cities, even in towns hav- 3.1-3.5 39 27 1 ing nearly or slightly more than 1,000 people, which were the & 0.499 highest frequency sizes of towns. Population size was scaled as logs to base 10. After Long and Thiess. 3.6-4.0 32 17 1 1 & 0.495 Log size Number Number bats Bats/log size 4.1-4.5 27 10 2 1 1 towns bats & 0.505 <2.0 ~ 100 10 — 4.6-5.0 65 13 3 3 0 2 2.1-2.25 48 14 6.2 & 0.403 2.26-3.0 218 24 8.0 3.1 180 67 19.1 5.1-5.5 19 17 2 0 3.6-4.0 65 51 12.8 & 0.547 4.1-4.5 50 41 9.1 >5.6 4 1 1 2 1 4.6-5.0 10 86 17.2 & 0.717 5.1-5.5 2 38 6.9 5.6-6.0 1 9 1.6 Totals 215 104 13 7 2 2

Table Chi. 5. Relative abundance (RA) and species diversity of bats netted in southwestern Wisconsin, by habitat. Netting hours per habitat in parentheses. RA = number of bats/ number of hours netting. After Ainslie, 1983.

Northern Northern Southern Southern Southern Southern forest forest forest forest forest forest dry-mesic (5.8) mesic (6.2) xeric (17.5) lowland (20.8) dry-mesic (14.7) mesic (29.7) RA N RA N RA N RA N RA N RA N Little Brown Bat 129.3 75 104.8 65 109.1 191 88.0 183 84.4 124 68.0 202 Red Bat 17.2 10 11.3 7 0.6 1 1.9 4 0.7 1 11.1 33 Keen’s Myotia 3.4 2 17.7 11 5.7 10 2.4 5 4.8 7 3.0 9 Big Brown Bat 20.7 12 4.8 3 1.1 2 0.0 0.7 1 2.7 8 Georgian Pipistrelle 12.1 7 0.0 1.1 2 0.5 1 0.0 0.3 1 Hoary Bat 5.2 3 0.0 1.7 3 1.0 2 0.0 0.3 1 Total 187.9 109 138.7 86 119.4 209 93.8 195 90.5 133 85.5 254 Species Diversity (H) .456 .341 .184 .089 .133 .295

TAXONOMIC ACCOUNTS / ORDER CHIROPTERA 141 ± .23 .17 ±.09 6.85 4.36 5.08 7.33 5.31 4.89 5 19.52 10.0719.4 7.03 10.01 5.55 5.98 6.93 –––– nd exponent value is N for some samples. 3 14.4 6.62 g 14.37 5 2 ± .8 ± .31 ± 8 6 5 1 34 4 – 13.1 7.0 4.15 4.76 8.69 37.3 4.17 9.3g 13.53 7.69 4.48 5.63 – 38.0 6.67 9.0 15.0 54.17 6.83 24.7g 17.71 10.16 6.06 6.78 37.5 9.3 15.0 35.5 8.0 – 14.88 7.48 5.59 4.98 ± 8± ± 6.9 4.3 ± ± 5.3 ± 2.2 ± 4.6 ± 2.1 ± 1.1.4 ± ± .44 ± .18 ± .25 ± .14 ± .14 ± .24 ± .14 79.8 37.0 7.31 10.54 33.3 4.38 6.3 13.13 ± 2.8± ±2.6 ±1. .97 ± 1. ± ±96 6-6.5 .14 ± .1 ± .22 ± .34 ± ± .5 ± 2.8 ± .8 ± .5 ± 1.0 ± .98 ± .9 ± .48 ± .15 ± .3 ± .1 length length length length length length l. Skull br. t-r depth ± 4.11 ± 2.5.8 ± ± 1.3 ± 1.2 ± .34 ± .25 ± .18 ± .41 88 40 9 15 36 7 – 15.32 7.50 5.9 5.7 ± 3.16 ± 3.1 ± .150 ± 110 .58 ± 44 10.7 14.3 45 Bayfield Co. 3.0 ± ± 1.9 ± .69 ± .76 ± 1.20 ± .35 ± ± .34 +/- .29 Missouri 91 38 10 12 – 8 – 14.1 8.3 5.1 5.1 FF Limery C. Maiden Rock 86 85 35 35 6 10 10 10 33 4.5 – 12.6 6.8 4.05 5.0 F Sauk Co. 102 46 6 9 39 3 10.5 14.2 7.9 4.9 5.7 M’s8 ± 3.2 ± ± 1.1 ± 1.26 ± 2.6 ± .98 ± 2.25.5 ± ± .23 ± .26 ± .99 7 F 110.2 40 13 16 47 10 M’s Beetown C. 3 F Beetown C. 79 34 9 11 34 4 – 12.95 6.8 4.1 5.3 5 M’s2 Sauk Co. F 90.7 Sauk Co. 42.3 75 6.67 39 7 8 37 4 8.5 6 F’’s & Grant & Sauk C. 126.7 57.0 8 M 87331015377––––– 1F“ 89351013387––––– 1F“ Drummond,6 87.6 M’s & F’s Indiana 34.6 89.7 10.14 35.17 14.71 9.08 38.1 7.0 – 14.93 7.59 – 5.02 4 M2 F Grant Co. 86.0 “ 89 38 9 16 37 7 – 15.5 7.85 6.0 4.7 1 F2 embs 1 F11 M’s Brown Co. Pt. Vic.Stevens 102 42 8 11 42 3 – 16.0 7.5 – 5.0 External and cranial measurements for Wisconsin’s species of bats. Means ± standard deviation. Sample size is N, a Table Chi-6. Table Chi-6. Table Chi-6. Table Chi-6. Table Chi-6.

Perimyotis subflavus Perimyotis Lasiurus borealis Lasiurus Lasiurus cinereus Lasiurus SpeciesMyotis lucifugus N Sex LocalityMyotis sodalis Total Tail Hindfoot Ear FA tr. Weight Greatest Cranial Max Cranial Myotis keenii Nycticeius humeralis Lasionycteris Eptesicus fuscus

142 THE WILD MAMMALS OF WISCONSIN Order LAGOMORPHA Pikas, Rabbits, and Hares

The ancestry of the lagomorphs is not known for certain; insectivores and primates have been suggested. Owing to their mutual gnawing and destructive habits, the lagomorphs long had been confused with the rodents, until Gidley (1912) separated the two orders, and in some quarters they are confused today by Skull of a lagomorph. Anonymous artist. Note skull lay people. These two orders of mammals fenestrae, upper incisors, and diastema of the jaws. actually have maintained separate lineages at least as early as their first preserved fossils— tail is always short with a conspicuous bunch in the early Eocene Epoch (perhaps late Pale- of fluffy hairs. The hind foot may or may not ocene, see Meng et al., 1994) for the rodents, possess a dew claw (vestigial toe nail), and and the late Paleocene for the ancient Lago- the other four hind toes (beneath a dense hairy morphs. Lagomorpha resemble rodents in pad) are webbed for support (holding the toes having two chisel-like upper incisors for gnaw- together while leaping). The forefoot has five ing. Unlike rodents, the pikas, rabbits, and toes. The testes are extruded into a scrotal hares have a second pair of upper incisors, sac, but lie under the skin situated a little an- tiny and peg-like (or spicule-like) sited imme- terior of the conical penis. There is no bacu- diately behind the first pair. The chisel-like lum as seen in rodents. incisors and gnawing behavior have evolved For its herbivorous diet, the molars are com- in parallel. Modern Lagomorpha are divided prised of transverse loph-like crests, and there is into two families, the Holarctic pikas (Ochoto- a pronounced gap (diastema) between the chew- nidae) and the cosmopolitan rabbits and hares ing teeth and the chisel-like incisors extending (Leporidae). A secondarily derived (not prim- most of the length of the rostrum and beneath, itive) cloaca is seen in the pikas (Long, 1970). also of the dentary, i.e., the premaxillaries, max- There are rabbits and hares in Wisconsin. illaries and nasals. The zygomatic arch projects slightly, both anteriorly and posteriorly, as orbital processes, and a frontal (supraorbital) exten- Family LEPORIDAE Gray sion partially enclosing the orbit above has similar anterior and posterior processes. The poste- “Naturalists have sought... to distinguish rior process may come in close proximity to the between the Rabbits and the Hares; but braincase (in Sylvilagus). At various places on although the young of the former are blind and the skull surface, the bone is exceedingly porous naked when born... and the latter are clothed or perforate, and this screen-like structure is with hair, and have the eyes open, and in their termed fenestrate (meaning “with windows”). habits there is this difference, that the Rabbits There is a clavicle, often lacking in leaping ani- burrow, whilst the Hares make a “form”... on mals. The tibia and fibula are fused distally. The which they squat.” – G. R. Waterhouse, 1848, skin is very thin and emits water for evaporation Natural History of the Mammalia, Vol. 2. by insensitive perspiration. Lagomorphs often reingest their own feces (coprophagy) for more The Leporidae, or leapers, are charac- efficient digestion of vegetal carbon compounds. terized by their diminutive second upper inci- Rabbits differ from jack rabbits and other hares sors, and by elongate hind limbs used for hop- (genus Lepus) in giving birth to naked, blind, ping and leaping (saltatorial locomotion). The helpless (“altricial”) young in a nest made of the

TAXONOMIC ACCOUNTS / ORDER LAGOMORPHA 143 mother’s ventral hair. In Lepus there is no nest 3 Feet relatively large for standing on snow, at all, and the young are born on the ground. size medium (total length usually exceeds Their eyes are open at birth, the body haired, 450 mm, hind foot 110-150 mm), dor- and the active young are said to be “precocial.” sal fur in summer dark or pale grizzled In Lepus americanus and L. townsendii the fur ochraceous and creamy brown without usually molts to white in early winter, to brown- orange rust on the nape, tail dusky above, ish in spring and summer. white below, in winter usually pure white (with pale buffy gray underfur entirely Key to the species of hidden) except for black eyelids and ear- Leporidae in Wisconsin tips ...... Snowshoe hare Lepus americanus 1. Size small (total length 460 mm or less, 3’ Feet relatively small but correlated with hind foot less than 100 mm), orange on large size (total length exceeds 550 mm, nape,skull usually less than 75 mm in hind foot about 145-172 mm), dorsal fur length, interparietal distinct from parietal in summer buffy or creamy gray and bones posteriorly, postorbital process of slightly grizzled, no markings on the neck, frontal bone fused to braincase (to cranial tail all white, in winter the fur is pure frontal) instead of flaring away from the white excepting black ear tips ...... braincase, jugal of zygoma narrow and ...... White-tailed jack rabbit hardly tapered, and interpterygoid fossa Lepus townsendii (depression or cleft between the pterygoids) narrow ...... Eastern cottontail Sylvilagus floridanus Genus Lepus Linnaeus 1’ Size large, skull elongate, interparietal Jack Rabbits and Hares fused posteriorly, postorbital process of frontal flares away from the braincase, See account of Order Lagomorpha for distin- jugal expanded in the middle and taper- guishing features of jack rabbits and hares ing at either end, interpterygoid fossa from those of cottontail rabbits. wide ...... 2 2. Summer and winter pelage on the back grizzled grayish to yellowish brown, whit- Lepus townsendii Bachman ish at hair bases, grayer on cheeks and White-tailed Jack Rabbit flanks, nape and feet buffy, tail blackish above, zygomatic arches broad anterior- Lepus townsendii campanius Hollister ly, nasals exceptionally wide and exceeding 42 mm in length ..... European hare 1837. Lepus campestris Bachman. J. Acad. Nat. Sci. Lepus capensis Philadelphia, 7:349. Not Lepus cuniculus (might occur in Wisconsin, introduced campestris Meyer, 1790. Type from onto Upper Peninsula of Michigan). Saskatchewan, near Carlton House. 2’ Fur in summer grizzled dark brown or yel- 1915. Lepus townsendii campanius Hollister. Proc. lowish tan, hairs not whitish basally, nape Biol. Soc. Washington, 28: 70, a renaming of never orange or yellowish, feet brown or L. campestris. tan, zygomata broader across posterior processes than across the anterior process- The name townsendii honors J. K. es, nasals not as broad, though relatively Townsend, who collected the holotype of the wide in comparison to Sylvilagus, but al- nominate race at Walla Walla, Washington. ways less than 42mm in length ...... 3 Lepus means leaper.

144 THE WILD MAMMALS OF WISCONSIN Description. The white-tailed jack rabbit from Minnesota as a game animal, and persist- is readily identified by large size and white ed for years in widely separated marshes and tail, as well as by its prairie surroundings. The prairies. The range expanded from 1944 until skull is less massive than in the European hare 1953 (Lemke, 1956) but now has greatly di- L. capensis (see Key above), and the length minished. The species may be eradicated. No of the nasals is less than 42 mm. The postor- one I have consulted in the Department of Nat- bital and antorbital process flare away from ural Resources verifies that any remain. It closely the skull. Chromosomes are 2N = 48 (Hsu resembles snowshoe hares in winter. I have one and Benirschke, 1971). specimen, from northern Langlade County. The color in summer is a creamy or buffy Status. The white-tailed jack rabbit seems gray intermixed with fine brownish hairs, and to have vanished from Wisconsin, and the rea- the ears are tipped with black. The underparts sons are unknown. It quietly disappeared in are white as is the tail. In winter the fur is white, the last 30 years. None of several reports of except the black tips of the ears, but the fur its occurrence could be verified. The preferred and the forefeet are suffused or faintly washed habitats of open marshland and prairie having with buff. Occasionally a thin black line is seen wide horizons, i.e., grasslands of vast expans- on the upper side of the fluffy tail. Young, hav- es, are also fast vanishing and almost gone. ing fewer guard hairs, show more underfur The last known jack rabbit in central Wiscon- color (Lim, 1987). The winter molt occurs in sin (Buena Vista Marsh) was collected in 1966, November and early December (Lim, 1987). by one of my undergraduate students, who The hind feet and ears grow faster than donated it to the collection in Madison. Will- other body dimensions (Lim, 1987). Lengths iam Smith (personal comm.) reported that in vary to 27 inches, and a large hare may weight the late1960’s a few existed on the Military 10 pounds ( = 4.5 kg). Ridge prairie country of Grant and Iowa coun- Dental Formula. I 2/ 1, C 0/0, P 3/2, ties (southwest Wisconsin). More than a third M 3/3 = 28. of the 19,000 jack rabbits killed in 1951-52, Geographic Range. Possibly this huge, came from Barron, Clark, Eau Claire, Mara- white-tailed jack rabbit was a rare but natural thon, Portage, Waushara, and Wood counties. denizen of southwest prairies in Wisconsin (Co- There are no such numbers today, although rey, 1912), although Jackson (1961) did not the species is secretive (but often seen, if think so. Long (1970) listed it as possibly an present, on plowed fields where cover is ab- introduced species. It was widely introduced sent). A few ranchers on the Buena Vista Marsh claim the species is present there, but more likely they see snowshoe hares. Habitats. The white-tailed jack rabbit inhabits open grasslands, prairies or marshes. Occasionally it ranges onto plowed and mowed fields. The jack rabbit may prefer one place on which to rest, called a form, where the vegetation is pressed down. A form is about one by two feet in dimensions. In winter this hare may scratch out a depression in the snow, even digging sometimes a hole for itself in the snow. Foods. Lepus townsendii feeds on green vegetation when available, and buds and bark Skull of Lepus townsendii. Dorsal, ventral views. when greens are not available (Jackson, 1961).

TAXONOMIC ACCOUNTS / ORDER LAGOMORPHA 145 Maps showing geographic distribution of Lepus townsendii in Wisconsin and North America. The patchy distribution in Wisconsin after Domke, 1973. Most recent occurrences are shown by dots.

Reproduction. Breeding commences in as high as 71 (Kline, 1963). Usually the jack April or May, with a gestation of about one rabbits assemble in groups of two to four and month. The precocial young may be brought go hopping over the prairie in the darkness. forth in a form or dropped on the ground. They are seldom seen in the day time, unless There seems to be a single litter in the year. on plowed ground or the roadway. Litter size is about four (3-6). Weaning may Geographic Variation. None was evident not occur until the young “leveret” is eat- in Wisconsin. ing green food, at about 4 weeks of age or Specimens examined. Total, 3. Portage even later. Co.: Buena Vista Marsh (UW20425, 22 Mortality. Hawks, owls, and occasional- March 1969). Langlade Co.: 5 mi. N Antigo, ly coyotes kill this jack rabbit, and cars often near Bryant (No. 1021), 24 Oct. 1966. Door kill them on the highways. Once thousands Co. Approaching Sturgeon Bay Sight record. were harvested in Wisconsin by hunters 1966. Sheboygan Co. Holland 1 UW Wild. (1951-1952, also see Scott, 1947). Lice, Ecol. [Local. not found.] fleas, probably ticks, tapeworms, and probably roundworms parasitize this hare. The species can carry tularemia (Jackson, 1961) and Lepus americanus Erxleben some other diseases (Lim, 1987). Home Range and Density. The home 1777. [Lepus] americanus Erxleben. Systema regne range is given by Seton (1953) and Jackson animalis . . . . 1: 330. Type from Hudson Bay, (1961) as 2-3 km in diameter. Jackson (1961) Canada. reported the kill of jack rabbits in l951-1952, as l9,383 animals. The populations fluctuat- “ ... a specimen of the L. americanus ... November, a ed, but today certainly nothing like those num- time in which the fur is undergoing its change from bers of hares remains in Wisconsin, where summer to winter colouring. white, with the exception the species was once abundant. Little is known of the head, cheek, back, and chest ... On the back, about density, reported as 3-9 per km2, but the coloured hairs are considerably longer than on the

146 THE WILD MAMMALS OF WISCONSIN sides of the body, where they are white... a consider- white. There are two molts per year. Hares able portion of the summer hairs have been shed from in this collection are seen in molt in Novem- the back white hairs... Were the long and scanty ber and December, and by late April the coloured hairs to be plucked... it would present the white is nearly replaced. same condition of full winter pelage... the longer and The Total length usually varies from 14- coarser hairs are shed and replaced by equally long, 20 inches (360-500 mm); weights vary up to white-pointed hairs... the dense under fur... becomes four pounds ( = 1.8 kg). A large male from longer.” — G. B. Waterhouse, 1842, The Natural Langlade and four females from Portage coun- History of Mammals, Vol. 2. ties are, respectively, as follows: Total length 630, 436±22; length hind foot 145, 135±7;ear length 72, 74.3± 5; greatest length Lepus americanus phaeonotus J.A. Allen skull 79.1, 77.3± 3.4; zygomatic breadth 39.2, Snowshoe Hare 38.9± 1.1; length of nasals 33.5, 31.8± 2.6; maxillary toothrow 15.4, 15.0± 0.8. 1899. Lepus americanus phaeonotus J.A. Allen. Dental Formula. I 2/ 1, C 0/0, P 3/2, Bull. Amer. Mus. Nat. Hist., 12:11. Type lo- M 3/3 = 28. cality: Hallock, Minnesota. Geographic Distribution. The range of this species has contracted since the turn of The scientific name means American the century (Jackson, 1961). The same thing leaper or American hare. “Snowshoe” refers has happened in lower Michigan (Baker, to the huge hind feet, which better enable 1983), where the hare occurs chiefly in the the hare to hop across snow (the chief force northern counties, and throughout the Up- of leaping is focused on the hind feet). The per Peninsula (see Maps). hare also was called a snowshoe rabbit, or Status. Numbers of this hare in Wiscon- the varying hare. sin have not changed appreciably since Jack- Description. See Key to the Leporids son’s (1961) mapping. As a game animal the above. Not so large as Lepus townsendii species is hunted but also managed, or in this or L. capensis, and resembling the Eastern case protected at least by hunting season and Cottontail Sylvilagus except that the ears possession limits. and especially the hind feet are larger (and Habitats. The snowshoe hare inhabits heavily padded with bristly hairs). The skull the northern boreal communities, conifer differs from that in Sylvilagus in its flaring swamps where tamaracks and cedars thrive, postorbital processes, and in partial fusion spruce-fir woodlands, and wetlands such as of the interparietal, and from other Lepus alder swamps, aspen-birch marshlands, and by the rudimentary anterior process on the cat-tail marshes (Grange, 1932; Jackson, supraorbital extensions of the frontal. There 1961; Baker, 1983). are six mammae. Ground cover of vegetation is essential The summer pelage of this hare on the for this hare. It prefers edge habitats. Some upper parts is ochraceous brownish gray, of- prime habitats are thickets of willow Salix, ten a deep ochraceous brown with intermix- alder Alnus, and aspen saplings Populus (Kei- ture of black (especially in young hares). th et al., 1993; Keith and Bloomer, 1993). There is no rusty orange nape patch as in This hare rests in scratched out forms, Sylvilagus or domestic Oryctolagus. The un- under sheltering grass, bushes, or a fallen log. derparts are white. The ears are tipped with Occasionally it burrows under ground or into black. The winter pelage is all white except snow banks, but not often (Baker, 1983). eyelids and ear tips. Molting individuals, of Foods. The snowshoe hare feeds on course, show various patterns of brown and green vegetation, and winter buds, twigs, bark,

TAXONOMIC ACCOUNTS / ORDER LAGOMORPHA 147 and conifer needles. The fecal pellets are of- Young are born by mid-April (usually 3, 1-7). ten eaten vegetation, and winter buds, twigs, The young are furred and their eyes open. They bark, and conifer needles. The fecal pellets weigh up to 25 ounces. Females do not breed are often eaten (coprophagy) to attain maxi- the first year. Older females mate repeatedly; mal food value from complex carbon com- they may bear three or four litters per season. pounds in plants (Bookout, 1959). Bookout Later litters average larger in litter size (Rowan further suggests (1965) the best winter for- and Keith, 1956). When the population cycle is age is yellow and paper birch, white cedar, on the rise, the mean litter size may increase. sugar maple, trembling aspen, American elm, Each neonate is furred with eyes open, and jack pine, and red pine. Other plants used as weighs about 65-80 g. Neonates can crawl forage are white and black spruce, hemlock, about, but rather feebly. By nine days they have which pine, and beaked hazel. doubled in weight and begin to eat green vege- Reproduction. The snowshoe hares mate tation. By three weeks of age or a little later the promiscuously. Occasionally the leverets may leverets are weaned and leave the form. Baker attempt to breed (Keith and Meslow (1967). (1983) reports the young “freeze” (that is, re- Mating begins in March (when hares are sup- main motionless) in time of danger, and the posed to be “mad” because the males may be mother hare “distracts adversaries away from seen leaping erratically about) and lasts until early her litter.” The average productivity for Michi- September. Gestation is about 35-36 days. gan hares is 6.5 young per female (Bookout, 1965). Longevity is 3-4 years, but there is a report of 8 (Crandall, 1964). Mortality. The snowshoe hare is an important forage species in the North Woods, prey for most of the carnivores and birds of prey with which it co-exists. In fact, high frequencies of hares increase the numbers of predators, and lows decrease them. In central Wisconsin, coyotes ate most of the hares radio-collared by Keith et al. Home Range and Density. Home range varies from 5-57 acres (2-22.8 ha). Movements are extensive in summer. Breeding males have larger territories than other hares. In central Wisconsin, in 1990-91, maximum home ranges varied from 1.6 to 0.8 ha in prime habitat of willow, alder and poplar. In patchy environments of willow, alder and poplar, densities varied from 1.0 to 2.7 hares per hectare, and over the total range from 0.3 to 1.3 / ha (Kei- th et al., 1993; Keith and Bloomer, 1993). Dispersals were most frequently made from small patches. Unless the patch was large (>5 /ha) or the number of snowshoe hares large enough (10 or more) the hares were likely to Skull of Lepus americanus. Dorsal and ventral views. vanish. The population was declining, and the Juveniles are compared below, as their different proportions chief predator was the coyote (Keith et al., resemble cottontails. 1993; Keith and Bloomer, 1993).

148 THE WILD MAMMALS OF WISCONSIN Besides annual fluctuations there are multi-annual population fluctuations, with population peaks every 9-11 years (Keith and Wind- berg, 1978). The cycle also has been referred to as the 9-10 year cycle and the ten-year cycle. Some high peaks occur every six years and others as seldom as every 13 years. Some relevant studies among a great many include Adams (1959), Adamsic and Keith, 1978, Adamsic et al., 1979, Arnold (1956), Book- out, (1965), Brand and Keith (1979), Brand et al. (1975, 1976), de Vos (1951), Errington et al. (1940), Keith (1974), Keith and Windberg (1978), MacLulich (1937), Meslow and Keith

Maps of distribution of Lepus americanus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER LAGOMORPHA 149 Table Lag-1. Suspected Fluctuations in Multi-annual Jordan Swamp, central Wisconsin. After it Cycles in Mammals that occur in Wisconsin. After Dymond hopped up to me fearlessly, I picked it up and (1947), with Pitymys ochrogaster added as a possibly cyclic petted it. After I put it down, it went hopping mammal. away without showing any concern. Species Cycle (Yrs.) and Range (Yrs.) Additional Natural History. Bittner and Lepus americanus 9-10 (=10) (8-11)* Rongstad (1982) wrote an excellent review Lynx canadensis 9-10 (=10) (7-12) of biology for Lepus americanus. Sciurus carolinensis approx. 5*** Jackson (1961) Geographic Variation. None has been Microtus pennsylvanicus 2-4** (1-4) observed in Wisconsin. Hares on Isle Royale Pitymys ochrogaster ?2-4 May be irregular in Lake Superior have been referred to L. a Ondatra zibethicus 9-10 (8-12)*** Vulpes vulpes 9-10 (8-13)*** americanus (Baker, 1983). Martes pennanti 9-10 (8-11)*** Specimens examined. Total, 43. Ash- Neovison vison 9-10 (6-12)*** land, Bayfield, Burnett, Chippewa, Door, * Range revised to 6-13, see account of Lepus americanus. Florence, Forest, Iron, Jackson, Juneau, ** Cycle is often 2-3 years, or 4-5, often irregular, and has been Langlade, Lincoln, Marathon, Marinette, recorded 4-5. Oconto, Oneida, Portage, Price, Rusk, Tay- *** All workers I consulted knew of no regular cycles in these lor, Washburn, Waushara counties. mammals in Wisconsin.

(1968), Pettingill (1976), Todd et al. (1981), Genus Sylvilagus Gray Windberg and Keith (1976), and Wood and Cottontails Monroe (1977). In Michigan, the kill by hunters varies approximately from 150,000 to See Description under Sylvilagus floridanus. 750,000 hares per year (Baker, 1983). The numbers of this species throughout “Bred and born in a briar patch! And with that he Canada regularlyfluctuate in 10-11 year cycles skipped away as lively as a cricket in the embers.” of abundance (see works of Lloyd Keith, of — Brer Rabbit to Brer Fox (by J. C. Harris, Stories the University of Wisconsin—Madison, and his of Uncle Remus) many associates), but this hare shows irregular fluctuation in Wisconsin. Perhaps the coincidental effects of predator-prey interactions and Sylvilagus floridanus (J.A. Allen) seasonality cycles of vegetation abundance are Eastern Cottontail affected by Wisconsin hunting harvests. W.E. Scott (1952) summarized Wiscon- 1890. Lepus sylvaticus floridanus J. A. Allen. Bull. sin harvests (see Jackson, 1961) from 1931- Amer. Mus. Nat. Hist., 3:160. 1949. From 1950 until 1982, the harvest numbers are listed by Wise (1989) as follows: Sylvilagus floridanus mearnsii (J. A. Allen) Year Harvest Year Harvest 1950 154,428 1980 392,800 1894. Lepus sylvaticus mearnsii J.A. Allen. Bull. 1960 61,629 1981 232,900 Amer. Mus. Nat. Hist., 6:171. Type from Fort 1972 101,700 1982 163,766 Snelling, Minnesota. 1979 404,000 1904. Sylvilagus (Sylvilagus) floridanus mearnsi: In the winter of 1999-2000, the harvest of Lyon. Smith. Misc. Coll., 45:336. snowshoes was estimated as 61,858, mostly from Price, Oneida, and Rusk counties (Dhuey, 2000). Sylvilagus floridanus means the Flori- Remark. Once, in early summer, I ob- da wood rabbit. Eastern cottontails occur served a young snowshoe hare deep in the mostly in the eastern United States.

150 THE WILD MAMMALS OF WISCONSIN Description. The cottontail is the smallest black, brown, and buff, and underfur char- of Wisconsin leporids (total length 460 mm or coal gray. The throat and legs are reddish less, hind foot less than 100 mm, skull usually brown. Although there is no seasonal color less than 75 mm in total length), with skull change to winter white, as in Wisconsin Lep- narrow, nasals tapered anteriorly, interparietal us, the cottontail molts twice a year, one molt distinct from adjacent bones, fenestrae lack- before winter and another in the spring. Each ing on rostrum, anterior processes of supraor- eye is ringed with a bold white circle. bital extension of frontal rudimentary, posteri- In spring and early summer eastern cot- or process fused to braincase, hind foot small, tontails in central Wisconsin, half-grown or jugal of zygoma slender. There are four pairs older, show high frequencies of an unde- of mammae. Diploid chromosomes are 42. scribed and fairly conspicuous white stripe The eastern cottontail shows grizzled pattern in the grizzled fur. The band or stripe brown upper parts in all seasons, rusty or- commences above the hind limb and extends ange or reddish brown patch on nape of neck, forward approximately 100-150 mm on ei- underparts whitish or grayish, tail short, ther side. The width of the band is approxi- brownish gray above and conspicuous white mately 10-20 mm. This pattern seems possi- fluff below. Guard hairs are coarse, variably bly associated with molt because the dorsum is fresh brown, and ventrally the belly fur is gray. The grayish white stripe interrupts the grizzled brown by a short grayish extension isolating on each flank a small patch of grizzled brown fur. The stripe also may be related to the folding of skin, where the limb bulges from the body wall. It is not a physical consequence of bulging, and it extends too far forward to be thus explained. The stripe is hardly apparent in autumn. The cottontail varies in total length from 16-18 inches (400-460 mm), and weighs up to 4 pounds ( = 1.8 kg). Females are larger Skull of Sylvilagus floridanus. Adult. Dorsal and ventral than males. Some measurements of eight males views. from Portage County are as follows: Total length 427± 55; hind foot 99.6± 6.5; length ear 60.32± 5.6; greatest length of skull 74.2± 1.5; zygomatic breadth 36.7± 1.5; length nasals 32.6± 1; maxillary toothrow 15.0± 0.7. Dental Formula. I 2/1, C 0/0, P 3/2, M 3/3 = 28. Geographic Range. Cottontails are widespread and abundant throughout Wisconsin and the Upper Peninsula of Michigan. Found on some islands as well (see Map). Status. In many years the eastern cottontail is very abundant, concentrations occurring Skull of Sylvilagus floridanus. Juvenile. Dorsal and in proximity to farms, residential areas, and even ventral views. Juveniles differ in proportion from adults, urban areas. The species is an important small- resembling snowshoe hares. game animal, a fur species (but of poor quality),

TAXONOMIC ACCOUNTS / ORDER LAGOMORPHA 151 a meat animal, and a prey species for carnivores, raptors, snakes, and other predators. It can be a pest in our fields, gardens, orchards, lawns, and cemeteries. Its presence is often made known by chewed saplings, gnawed naked of bark up to a height of 2-3 feet. This species followed the clearing of forests and settlements of the early settlers northward into the forests. Jackson (1961) reported that the species occurred as far north as Green Lake County in 1846 (Muir, 1913: 181). Nelson (1909) documents the arrival of cottontails in Douglas County in 1907, but it

Maps showing geographic distribution of Sylvilagus floridanus in Wisconsin and North America.

152 THE WILD MAMMALS OF WISCONSIN was reportedly common just south of there. counters followed by jumping, and a variety of Jackson (1961) found them in the Apostle postures likely associated with sexual odors Islands in 1919. Long (1978) observed them (Chapman et al., 1980). Gestation lasts 29-30 on Washington Island, in lawns and fields, days. Rongstad (1969) described prenatal de- never in the forests or swamps. velopment. The female builds the nest (see Habitats. Cottontails are found at the Home above), and the number born average edge of woods and fields, in lawns, shrubby about 5 (range 4-7) in Michigan, in Illinois 4- areas, brush piles, brambles and briars, thick- 6. One in central Wisconsin had nine (see beets and hedges, usually in or near stands of low). A female may produce 20-25 young in a deciduous trees. Low shrubs with dense season. Furthermore, the young born early in branches and thorny hedge are good cover the season are 12-23 percent of all cottontails types for rabbits. Also they are seen in grassy present in the fall. Young of the year also breed. fields and savannas, roadsides, disturbed ar- A life table by Lord (1958) began with a theo- eas, and cut-over forest. They are not so com- retical population of 1,000, which peaked at mon in corn and soybean fields. They are re- 5,688 adults and young by August. placed northward in boreal and wet habitats The blind, naked young weigh about 30 by snowshoe hares. g each, and are suckled twice a day. The eyes The eastern cottontail rabbit uses burrows open about the sixth day. In two weeks the in which to hide or stay warm, especially in win- furred young are out of the nest, although ter. It often appropriates burrows made by the mother offers them milk for some days. woodchucks and badgers. It also finds shelter in Some 25 years ago, on the south side of thawed and grassy openings in snow banks, in Stevens Point, I observed two or three small dense thickets and junipers, woodpiles, and in cottontails out of their nest reaching up to brushpiles. In the breeding season the female’s nurse at their mother’s breast. On 25 May nest is a small pit probably excavated, lined with 2003. Claudine Long observed in late after- fur plucked from her breast, and covered over noon in Stevens Point, an odd “pyramid-ap- with grass, leaves, or pine needles. pearance” resembling a rabbit. When she Foods. In Wisconsin, green vegetation, approached to approximately 8 m (measured seeds, flowers, bark of shrubs, oak and other later) a large cottontail bolted away. What saplings, even blackberry, fruit trees and fruits, caused the pyramid shape was a litter of young buds and twigs, herbs, grasses, clover, and (each approximately 120 mm in length). The cultivated crops (Allen, 1939; Hickie, 1940; young had been standing simultaneously at Haugen, 1942; personal observations) are the mother’s breast trying to nurse. When she eaten. In the caecum, a sac at the junction of left them, they crouched flat on the ground, the small and large intestines, where micro- bodies crowded together randomly, in a space organisms are prolific, complex carbohydrates about 0.6 m2. All were watching, but none are fermented and formed into soft, green moved. She counted nine young. When she fecal pellets. The large, green pellets are bent down they scattered. passed and re-ingested for further nutrition, Eye lens weights of the eastern cotton- a behaviorism called “coprophagy” (Geis, tail vary with increasing age. Lord (1959) plot- 1957; Kirkpatrick, 1960; Bailey, 1969). ted out these weights (maximum wt 200 mg). Reproduction. The male cottontail’s tes- After one year of growth the curve levels off. tes enlarge in Michigan from late February to Hoffmeister and Zimmerman (1967) found mid-March, and breeding continues until Sep- that at 5.5 months cottontails attain maximal tember. Lemke (1957) found one pregnant size. Petrides (1951) described fusion of the female on 21 November. The male gambols epiphyses of the radius and ulna between 7 about in a courtship dance, head-to-head en- and 18 months.

TAXONOMIC ACCOUNTS / ORDER LAGOMORPHA 153 Mortality. In Michigan Hickie (1940) esti- and at least nine kinds of roundworms (Jack- mated that in 1938, more than 300,000 hunt- son, 1961; Chapman et al., 1980). Young rab- ers bagged 2,000,000 cottontails. In 1974, bits often drown or chill and die in the nest. some 410,000 hunters killed an estimated 2.25 They are killed by automobiles and cats. One million, and in 1975, the take was 2.47 mil- may add to natural mortality the game harvest, lion (Baker, 1983). In Wisconsin, the numbers for this is the most commonly killed animal tak- are comparable. In 1932 (2,474,125), 1942 en by hunters, throughout the United States. (1,397,308), and 1954 (1,311,392) the re- Home Range and Density. Trent and sults compare closely. Low years dropped to Rongstad (1974) radio-tagged cottontails, only 382,186, which is still a phenomenal finding the males to wander in home ranges number for game animals. In 1999-2000, the about 10 acres (4 ha), diminishing to 4 acres harvest of cottontails was estimated at in the late summer (after breeding). Adult fe- 250,727, mostly from Grant, Dodge and She- males averaged 4.3 acres (1.7 ha) in spring boygan counties (Dhuey, 2000). and 21 acres in late summer. Other findings The list of natural predators is lengthy (Jack- on populations of cottontails have been ob- son, 1961, Baker, 1983, Chapman et al., 1980) tained by Hickie (1940), McCabe (1943), and for this important forage species, including coy- Lord (1963). The turnover rate of cottontails otes, foxes, domestic dogs, wolves, bears, prob- is 80 percent. ably raccoons and probably opossums, mustel- There is evidence for and against regular ids except otter, bobcat, domestic cats, hawks periodic cycles, but certainly there are highs and owls, crows, red squirrel, great blue heron, and lows of abundance. At my home in cen- and several species of snakes. Even small pred- tral Wisconsin, in jack pine-oak savanna, when ators can kill baby cottontails. the populations of cottontails were high (win- Cottontail rabbits harbor several deadly ter, 1991) my wife and I removed 41 cotton- diseases of humans such as tularemia, Pas- tails from one acre without any apparent ef- teurella tularensis, transmitted from rabbit fect on their numbers. Other cottontails moved blood (during cleaning of the rabbit for food). into the area to replace those removed. In three More than 10 human cases a year are con- years (1997-1999) not a single cottontail was tracted in Wisconsin. It may be passed from observed on this acre. In spring (2002-2003) rabbit to human or rabbit to rabbit by numer- we have seen several, but again in winters ous arthropods, including ticks, flies, mites, 2000-2003 not one. Because of the behavior fleas, even mosquitoes. Yellow specks on the of the rabbits in the summer of 1999, and five liver and spleen indicate the presence of tula- found dead, we assumed the population had remia in a rabbit. The rabbit also has trouble suffered a tularemia epidemic (see Mortality moving about and develops lesions. Eradica- above). Also, urban deer removed much of the tion of huge populations of rabbits often oc- cover, e.g., dense thickets of blackberries. curs in nature by this disease. Salmonella Keith and Bloomer (1993) found cotton- (Youatt and Fay, 1968), Q fever, Rocky Moun- tails on snow to suffer much higher mortality tain spotted fever, and equine encephalitus from predation than did Lepus americanus also have been reported in cottontails. Sick- (89% to 18% vs 84% to 63%). Their central ness from the raccoon round worm affects Wisconsin study suggests that predation (pri- some rabbits. Some rabbits suffer from Her- marily coyote predation) may eradicate some pes (Schmidt et al., 1992; and others). cottontail populations in northern forests Parasites include bot warbles, ticks, espe- where snow cover is usually heavy. cially in Wisconsin the rabbit tick (C. F. Long, Additional Natural History. Chapman et personal observation), many species of fleas, al. (1980) is an excellent reference source for mites, a fluke, numerous kinds of tapeworms, the eastern cottontail.

154 THE WILD MAMMALS OF WISCONSIN ancient mountain beavers. Tree squirrels (Sci- urus sp.) have specialized for arboreal habits, but their teeth seem primitive (Black, 1963). The Norway rat (Rattus norvegicus) was important in decimating the human populations of Europe in Medieval times by spreading, with the aid of flea parasites, the “black plague.” This pestilence persists today even in America. These rats and the house mouse (Mus musculus) may transmit directly or in- Eastern cottontail. Charles Schwartz. directly the plague, tularemia, Rocky Moun- Geographic Variation. There is a single tain spotted fever, salmonella, Lyme disease, subspecies in Wisconsin and the Upper Pen- Hantavirus, and other diseases. They also insula of Michigan. destroy stored food, documents, books, and Specimens examined. Total, 109. Ad- other property in our homes. In Wisconsin, ams, Columbia, Crawford, Dane, Dodge, rodents, along with cottontails and nuisance Door, Green, Green Lake, Iron, Kenosha, deer, are among the most injurious of mam- Kewaunee, Juneau, Lincoln, Manitowoc, mals (causing over $1,000,000 damage an- Marathon, Marinette, Milwaukee, Oconto, nually). The muskrat Ondatra [historically the Oneida, Outagamie, Ozaukee, Polk, Por- most captured fur animal] is probably the most tage, Racine, Rusk, Sauk, Sheboygan, beneficial furbearer in the world. Trempealeau, Washington, Waukesha, Wau- What animals are more endearing than paca, Winnebago, Wood counties. gray and fox squirrels (Sciurus), the beautiful striped and alert little chipmunks (Tamias and Eutamias), and the other squirrels for that ORDER RODENTIA matter? The graceful and elegant forest deer Gnawing Mammals mouse (Peromyscus maniculatus maniculatus) is a lovely and gentle little mammal. Con- Almost half of the species of mammals in the sider the industry of the “busy beaver” (Cas- World (approximately 2,000 of 4,600 mam- tor canadensis), and its tremendous impact malian species) belong to the Rodentia. About on American destiny and history by virtue of a third of the approximately 1,000 mamma- its valuable fur luring westward (and into Wis- lian genera of the world are rodents. Not only consin) those early explorers and settlers from are there many species, but a great number the eastern colonies. have amazing fecundity. Not surprisingly, they Many rodents are beneficial to man: eat- show high relative abundance in most terres- ing weed seeds and insects, making soil (see trial habitats. Rodents first evolved in the late below), and planting forest trees (by burying Paleocene Epoch and showed spectacular seeds). Not fully appreciated is their absolute- success by the Eocene. Although those an- ly essential role as “prey species.” Most Car- cestral short-legged gnawers are extinct, there nivores, beneficial snakes, and birds of prey are some still with us (e.g., mountain beavers depend on rodents for food, in one season or Aplodontia) that have evolved very little in another, in one place or another, so that our most of their characters, i.e., retaining many ecosystems are rich not only in nutrients and primitive features. The marmots Marmota, potential energy, but providing many species from North America and eastern Eurasia, in- with an array of food sources. cluding Wisconsin’s woodchuck (Marmota The vole Microtus pennsylvanicus may monax), have progressed little beyond the make up to 40 percent or more of the animal

TAXONOMIC ACCOUNTS / ORDER RODENTIA 155 biomass in the marsh, where it causes little trees. Other rodents, which are not too dis- harm to humankind. In these communities, it similar from squirrels in muscle pattern, in- may comprise 90 % of the mammal biomass. clude pocket gophers Geomys bursarius. Golley (1960b) showed the importance of this The Myomorpha include the mice and rats vole in funneling sun energy caught in plants that feature a slip of the masseter muscle ex- to the carnivores that feed on the voles. The tending through the infraorbital canal, finding voles consumed 1.6 % of available plant en- a new “origin” on the rostrum of the skull well ergy, and weasels consumed 31 percent of anterior to the zygomatic arch. That special- energy stored in the voles. The abundance of ization and generally small size must have been this vole may allow raptors and Carnivores to adaptive, because no other mammals have attain high densities. prospered as has this group. Myomorpha is a Rodents differ from lagomorphs in pos- dominant form of mammalian life, if one mea- sessing only a single pair of upper incisors. sures dominance by abundance and relative There is a diastema and loss of canines, com- abundance. In Wisconsin, the harvest mouse parable to that in lagomorphs. Some premo- Reithrodontomys, deer mice Peromyscus, lars are retained in most rodents, but in many voles, bog lemming, and muskrat all show the mice there are none. The radius and ulna are myomorph pattern. The hystricomorph ro- not fused distally in Wisconsin rodents as in dents have the infraorbital (= antorbital) canal rabbits. There is usually a large caecum to greatly enlarged as a passage for an extensive facilitate digestion of plant carbohydrates. The portion of the masseter muscle. In Wisconsin rodents tend to be small, although the beaver the porcupine is the sole member of this group, sometimes exceeds 90 pounds. which apparently arose in South America. Hys- In the evolution of gnawing and chew- tricomorphs on that continent are diverse in ing, several peculiar patterns of muscle attach- number of families and genera. The Wiscon- ment appeared leading to success of some sin zapodid (“jumping”) mice also have an en- kinds of rodents and to one means of classi- larged infraorbital canal, and resemble Hystri- fying suborders of rodents. Without delving comorpha in this character. into the controversial taxonomy of higher In Wisconsin there are ten species of categories of rodents, there are several fun- squirrels (Sciuridae), the beaver (Castoridae), damental patterns necessary and used to pocket gopher (Geomyidae), harvest mouse broadly classify Wisconsin rodents. These and deer mice, four voles, the bog lemming, patterns are “sciuromorph”, “myomorph”, muskrat, non-native house mouse and Nor- and “hystricomorph”. The most primitive of way rat (all in the Muridae), two species of our rodents belong to the Sciuromorpha, char- jumping mice (Zapodidae), and the hystrico- acterized by the simple attachment of the mas- morph porcupine (Erethizontidae). seter (which muscle lifts the lower jaw) to the sturdy zygomatic arch (zygoma). This is the typical attachment seen in mammals besides Family SCIURIDAE Gray sciuromorphs, for example, in cats, dogs, and Squirrels many mammals. The marmot has an expan- sive bony plate anterior to the zygoma that is Squirrels, bright-eyed rodents of diurnal hab- somewhat sulcate and to which muscle attach- its (excepting the nocturnal and crepuscular es. In this respect, the marmot hardly differs flying squirrels Glaucomys), have well-haired from the beaver, which has no deeper sulca- or bushy tails, a bow-legged stance useful for tion proportionally, although often that con- climbing the trunks of trees, and a prominent dition is believed true, and of course the bea- clawed thumb (e.g., Spermophilus) or small ver is highly specialized for chewing bark and thumbnail. Wisconsin squirrels include the

156 THE WILD MAMMALS OF WISCONSIN woodchuck (= groundhog), two ground squir- vers, distinct interorbital notches anterior rels, two chipmunks, three tree squirrels, and to prominent postorbital processes ...... two flying squirrels...... Woodchuck (= Groundhog) There is incredible species stacking and Marmota monax niche partitioning of habitat in Wisconsin. For 3' Zygomatic breadth less than 58 mm, an- example, in Portage County all ten squirrels terior lower premolar lacks paraconulid, occur, and on an acre by McDill Pond, seven zygomatic plate never sulcate, interobit- species co-existed (Table Rod-1). al notches and postorbital processes minute or absent ...... 4 4 Zygomata converging anteriorly, pelage Key to genera and species of conspicuously spotted, i.e., dappled with Squirrels in Wisconsin obscure spots ...... Ground squirrels Spermophilus 1. Lateral gliding membrane extends be- 4A Dorsal fur boldly marked with dis- tween forelimb and hindlimb, tail dors- tinct lines and rows of spots, hind oventrally. flattened Flying squirrels ..... foot 41 mm length or less, greatest ...... Glaucomys length of skull 46 mm length or less 1a Skull or hind foot 37 mm in length 13-lined Ground Squirrel ...... or more, whitish ventral hair grayish Spermophilus tridecemlineatus at bases, venter dusky, even on the 4B Dorsal fur dappled with pale spots throat...... Northern Flying Squirrel on tan color, head gray, hind foot Glaucomys sabrinus or skull exceeds 50 mm length ..... 1b Skull or hind foot less than 37 mm ...... Franklin’s Ground Squirrel length, ventral hair usually pure Spermophilus franklinii white, especially on the throat...... 4' Zygomata nearly parallel, tail long and ...... Southern Flying Squirrel well-haired, pelage never spotted ...... 5 Glaucomys volans 5 Greatest length of skull never exceeds 50 1' Lateral membrane absent, extending be- mm, venter pure white, dorsal pelage tween forelimb and hind limb, tail when brown or chestnut brown, baculum spi- fluffed not remarkably flattened ...... 2 cule-like ...... Red squirrel 2 Antorbital (= infraorbital) canal not de- Tamiasciurus hudsonicus veloped, the foramen piercing a zygo- 5' Greatest length of skull in adults more matic plate of bone, head with promi- than 50, venter ochraceous or whitish nent stripes ...... Chipmunks tinged with brown or gray, baculum dis- 2A Dorsal stripes to base of tail, a spi- coid except its base ...... Sciurus 5’A cule premolar anterior to P4/ ...... 5’A Much orange in fur, especially on ...... Least Chipmunk venter and tail, tiny spicule (= P 3/) Eutamias minimus never present ...... Fox squirrel 2B. Dorsal stripes extend to brownish Sciurus niger rump patch and not to base of tail, 5’A’ No orange in tail or dorsal fur, usu- P3/ lacking ..... Eastern Chipmunk ally whitish but brownish or even Tamias striatus ochraceous may occur on venter, tail 2' Antorbital canal well developed, head hairs show many white guard hairs, never striped ...... 3 many squirrels are melanistic, usu- 3 Zygomatic breadth greater than 48 mm, ally a spicule (P3/ ) in each upper anterior lower premolar having para- cheek toothrow ...... Gray squirrel conulid, zygomatic plate sulcate as in bea- Sciurus carolinensis

TAXONOMIC ACCOUNTS / ORDER RODENTIA 157 Table Rod-1. Squirrel food resources in central Wiscon- Genus Marmota Blumenbach sin. All herbivores or herbivore-omnivores, in season they eat Woodchuck and Marmots plant buds, insects, fruits, and fungi in season. All reproduced in the study area, but Sciurus niger, an edge species, was observed only once. The marmots are the largest of the squirrels, attaining weights in some forms up to 16 Species/niche Staples Oddities Habitat pounds (7.5 kg). The genus is exceptionally Woodchuck* Leaves, Oak leaves Edge, primitive among rodents, as are the ancient Grazer-browser shoots open woods aplodontids (mountain beavers). Marmots E. chipmunk* Acorns, Thornapple, Edge, have five toes on each foot, except in the Si- Granivore- hazelnuts, greenbriar woods berian species the thumbnail is lacking. The frugivore fruits Red squirrel* Pine cones, Eggs, Conifers, thumbnail is a small hoof or nail, functional Conivore- seeds fruits hardwoods more in standing on rocks or holding food granivore than in excavation (Long and Captain, 1974, Gray squirrel* Acorns, Mushrooms Woods 1977). Therefore, it is misleading to refer to Granivore- nuts, these fossorial but ancient squirrels as ground omnivore seeds S. flying squirrel* Acorns Birds, eggs Oaks squirrels, except in the sense that the name Granivore definition may refer to habits and not rela- 13-lined ground Seeds, Fledgling Dry grassland tionships. Also primitive is the stance, which squirrel shoots, is sub-digitigrade approaching plantigrade, Grazer-omnivore insects, more so than usual in the other bow-legged berries and digitigrade Sciuridae. The skull is flattened * May feed in trees and broad, as in the aplodontids, but postorbital processes are prominent. There is a primitive pattern of premolars, namely p 2/1, and the upper ones are not vestigial or lacking (as in more advanced squirrels). Anal glands are present. The coarse fur and fossorial habits confuse some people into mistaking woodchucks for badgers. The burrows are often taken over by badgers, foxes, skunks, and cottontails (in winter).

Lateral view of woodchuck skull. Flower & Lydek- ker, 1891. Marmota monax (Linnaeus) Woodchuck or Groundhog

1758. [Mus] monax Linnaeus. Systema naturae, ed. 10. 1:60, Type locality Maryland. 1904. [Marmota] monax: Trouessart. Catalog. Mamm., Suppl., p. 344.

The name Marmota from Latin means mountain rat (mar = rat), and monax reportedly was derived from an Indian name; this was the the scientific name used by Linnae- us. Woodchuck comes from the Cree Indian Woodchuck at burrow. By A. R. Dunmoor. 1900. word otcheck. The woodchuck is also called

158 THE WILD MAMMALS OF WISCONSIN groundhog, Siffleur (French-Canadian), whis- smaller, especially the deciduous third upper tle pig, and marmot. Groundhog’s Day, cele- premolars (dP3/ ). The broad molars above brated 2 February annually (especially in Penn- and below are rather high crowned with deep sylvania) probably was based on European basins in the lower teeth. A prominent re- myths about hedgehogs or Eurasian badgers, entrant angle invades each of the molars on and coincided with Candelmas, an ancient the labial surfaces, the angle in each of the religious celebration linking a sunny day with upper molars enclosed by a V-shaped loph. a cold spring. The groundhog supposedly There is usually a low loph anterior to each emerges to check the weather. If it sees its V. There are rudimentary internal cheek shadow, there will be six more weeks of win- pouches. Chromosomes are 2N = 38, FN = ter. If cloudy, spring comes early. In Wiscon- 62; the Y chromosome being small and acro- sin, spring never comes early. centric (Hoffman and Nadler, 1968). The Description. The woodchuck is a stout, baculum is broad basally and tapers to a vari- short-legged squirrel of medium size (large for able tip (in females there is a small os clitori- the squirrel family, about two feet in length) dis). The eight mammae are in two pectoral, with coarse grizzled fur and a long, fairly bushy one abdominal, and one inguinal pairs. tail. The skull is broad and flattened, with wide The upper parts are dark grayish or red- zygomata and wide mastoid breadth. There dish brown, grizzled by pale buff tips on the is a low sagittal crest and usually in adult wood- hairs. Basally the hairs are dark blackish or chucks a prominent lambdoid crest. In young- gray-brown, subterminally or terminally the er woodchucks the sagittal crest is undevel- hair is chestnut or ochraceous, and a con- oped and instead parallel sagittal ridges ap- spicuous buff tip is often present. The crown proach one another at the midline of the skull. of the head is black, often faded, and scatter- The occiput curves upward to the lambdoid ings of whitish buff hairs may be found around crests, and the rostrum curves downward an- the nosepad, extending onto the cheeks. A teriorly. The incisors are robust. The postor- little whitish may be seen over each eye. The bital processes of the frontals are prominent, feet are all blackish or dark drown, and the projecting laterally, whereas the postorbital bushy tail is nearly black. The black soles of processes of the jugal bones are low and arc- the feet are naked. The venter is variable in uate. The interorbital breath is rather narrow. tone, color and pattern. Usually it is some- The paroccipital processes are elongate. The what blackish but overlain with a tawny red two upper premolars are fairly well developed, and sometimes ochraceous or even buffy the milk teeth (present in young-of-the-year) ochraceous. Perhaps in the south there is less

Skull of Marmota monax. Dorsal, ventral views. Burrow of woodchuck. Missouri Conservation

TAXONOMIC ACCOUNTS / ORDER RODENTIA 159 black in the ventral fur. There is a single molt beginning in June lasting until August. Some melanistic woodchucks may be expected in Wisconsin (Jackson, 1961). Males are slightly larger than females in woodchucks (Jackson, 1961; Snyder et al., 1961) and the animals continue growing for several years. Some Wisconsin measurements are listed in the subspecies accounts. The following observed ranges in adults (in millimeters and kg) were given by Lee and Funder- burg (1982) for woodchucks from North

Maps showing geographic distribution of Marmota monax in Wisconsin and North America. The subspecies boundary follows Jackson, 1961.

160 THE WILD MAMMALS OF WISCONSIN America: Total length, 418-665; tail length, farther behind with the back feet. Usually a 100-155; hind foot length, 66-88; body mound of dirt is found in front of the bur- weight, 2.3-5.4. row’s entrance. If the animal opens a “plunge Dental Formula. I 1/1, C 0/0, P 2/1, M hole” or escape hole from within the burrow, 3/3= 22. there is no dirt pile there and the entrance to Geographic Range. Found in suitable it is seldom used. The main entrance is about habitats throughout Wisconsin except on is- 6-7 inches high by 7-8 inches wide (Jackson, lands in Lake Superior and Lake Michigan . 1961), sometimes larger. Usually there are This species prefers hills or embankments at at least two (sometimes three or four) entranc- the forest edge. es, and the burrow length may be 15-50 feet Status. Woodchucks are injurious when in length. The burrow usually is about two their den is near a vegetable garden, and they feet underground (1-5 feet), and there are occasionally damage a dike or embankment often tunnel branchings and a nesting cham- with their burrows. Personally, I do not ascribe ber. There is little vegetal material placed in to control methods mentioned by Jackson the nest, and one blind chamber may be used (1961). One may ask is gassing a rodent worse as a latrine (covered over fecal scats and dried than shooting it? If one has enjoyed this affec- urine). This home may be used, improved and tionate little rodent as a pet (Long and Long, enlarged by successive generations of wood- 1965), then one will object also to shooting it, chucks. like shooting a dog, and a lap dog at that. In The hibernaculum is similar to the nest any case, the woodchuck presently enjoys pro- chamber, but the burrow is plugged with soil tection in Wisconsin. If judged a nuisance the while the woodchuck sleeps. Occasionally a woodchuck may be trapped or killed. If one is skunk or rabbit may share the burrow sys- a pest, despite its aesthetic value, it can be easily tem, possibly unaware of its sleeping neigh- caught in a live trap and transported, hopeful- bor. Some diagrams of woodchuck burrows ly into a suitable habitat. Under legal protec- are depicted in books by Fisher, Jackson, and tion it seems that the woodchuck is improving Cory. A third type of home is more or less its status in the southern counties and increas- temporary, visited by mother and young, the ing also in the north. Although it is known to kind of subsidiary den called an “outlier” by have become scarce in some areas due to hunt- Long and Killingley (1983) and called “auxil- ing pressure, it is not certain that the wood- iary” by Kenneth Armitage (an expert on chuck was ever as widespread in Wisconsin as marmot behavior, and an alumnus of the Jackson (1961) assumed. In Illinois, Hoffmeis- University of Wisconsin). ter (1989) believed that when the settlers Foods. Except for many cultivated plants cleared the deciduous forests it made habitats from vegetable gardens, little is known about more suitable for the woodchuck. woodchuck foods. Pets I have raised loved to Habitat. Almost fossorial, the burrow- eat clover and dandelions, as well as a wide building woodchuck chooses for its den well- variety of foods we ate ourselves. Leaves from drained, sandy soils. The den is sited usually a sassafrass tree stuffed one wild woodchuck’s in open woods or the forest edge, where stomach, and I have seen them in trees in the woodland meets meadow, usually on a sunny spring eating oak leaves. Probably they are hillside or embankment, but occasionally near exceptionally hungry after a winter fast. One buildings, woodpiles, rocky outcrops, and had a full stomach of June beetles (Gianini, fencerows. Woodchucks require available suc- 1925), and another was attracted to pond culent vegetation, visibility, and sunshine. vegetation in Ontario (Fraser, 1979). Grizzell The burrow is excavated by tearing with (1955) examined 32 stomachs from individu- the foreclaws and teeth, and kicking the soil als in the spring, and found red clover, white

TAXONOMIC ACCOUNTS / ORDER RODENTIA 161 clover, grasses, chickweed, and alfalfa as fre- property would seem desirable to me. Dogs quently eaten foods. and automobiles are also major causes of Reproduction. Hoyt and Hoyt (1950) mortality, and if hunting of woodchucks were described copulation; a yearling male mount- legal the woodchuck probably could not hold ed a female, after “boxing” a few seconds with its own. Most road kills seem to occur after her. The female tucked her nose down and emergence from hibernation. Numerous held her tail up, while the male held her dor- young are killed by cars in late March, April sal fur with his teeth. The male copulated for and May. Some youngsters die in winter, and 8 minutes, then remounted for 5 minutes, others die if the burrow is flooded. Predation then again for 3. Males and females seem to seems of little importance in limiting wood- form pair bonds during the time of breeding, chucks. Foxes, coyotes, black bears, and bad- from late February through March in Mary- gers occasionally eat them (Grizzell, 1955; land, but in Missouri breeding begins a little Beule, 1949; Long and Killingley, 1983). earlier (Twitchell, 1939). As the time of par- Large snakes, hawks and owls occasionally turition, which has not been observed, the eat small woodchucks (Lee and Funderburg, female drives the male from her den. Gesta- 1982). Social stress reportedly can cause loss tion lasts about four weeks (Hamilton, 1934), of offspring (Grizzell, 1955). perhaps as long as 31-33 days (Grizzell, 1955; Occasionally woodchucks suffer parasit- Hoyt and Hoyt, 1950). Birth occurs in the ism, especially from arthropods such as ticks, period from early April into May. There are fleas, mites, and dipterans. A woodchuck may usually 4-6 young, occasionally as many as die from the infestation or from bacteria trans- nine. On average the number slightly exceeds mitted by the vectors (plague, perhaps spot- 4 young. Eleven newborn woodchucks yield- ted fever, tularemia). The roundworm Obe- ed 27.2 g average weight. They were nearly liscoides cuniculi has been reported from naked, toothless, blind and helpless. In a week woodchucks (Lee and Funderburg, 1982). the eyes opened, and the average weight for Home Range and Density. Unlike most each young at that time was 45-62 g (Griz- marmots, Marmota monax is less likely to zell, 1955). Davis (1964) observed young mingle together in colonies. On occasion 2-4 emerging from the burrows about 15 May, may be found sharing a den. Woodward each weighing about 300-450 g. By July the (1990) found high densities (up to 5.36 per young begin to disperse, usually to abandoned ha in fall) at highway interchanges (on the dens. They cannot be readily distinguished by man-made grassy roadsides). The roads ef- sight from the adults; their pelage is paler, fected linear shapes to the home ranges. The muzzle narrow, and their incisors lack brown home range seems considerably less than an pigmentation. Juveniles differ from adults in acre in central Wisconsin, although erratic small skulls, milk teeth, narrow rostrum, and wanderings have not been studied, and terri- smaller eye lens (Davis, 1964). Lactation re- toriality is not evident except around the ani- tards laying on of hibernation fat until wean- mal’s den. Movements are seldom over 100 ing. There is one litter born each year. Wood- m, and woodchucks from three dens I ob- chucks live between 4-10 years (Crandall, served were not observed to wander over 50 1964; Lee and Funderburg, 1982). m away. The smallest home range was of a Mortality. Even though the woodchuck yearling, killed two months after emergence is protected in Wisconsin, the animal is often by an automobile about 40 feet from the den. shot or trapped because of its forays to the This observed area was between standing garden. Some are eliminated because the den water and the road. is placed where people do not want it to be, Hamilton (1934) noted 30 occupied dens although seeing the young playing on one’s in less than 3 acres. In New York Manville

162 THE WILD MAMMALS OF WISCONSIN (1966) found 1/46 acres (18.4 ha), and Additional natural history. Swiecinski Twitchell (1939) found 1/36 acres (14.4 ha) (1998) reviewed the woodchuck’s natural his- in summer in Missouri. Grizzell (1955) count- tory. ed 18/100 acres (40 ha). Laundre (1975) Geographic Variation. According to estimated 30- 40/640 acres (256 / ha) on Jackson (1961) the southern woodchucks are the Upper Peninsula of Michigan. larger and less reddish tawny on the under- Remarks. Woodchucks are diurnal, al- parts. In the small sampling of Wisconsin skins though soon after emerging from hibernation in this collection, I saw none of the color dif- the hungry animals reportedly occasionally ferences described by Jackson (1961). There continue their activities into the night. Noctur- was no paler color evident in the south ex- nal activity is seldom observed (Grizzell, 1955). cept in older specimens showing some bleach- They are true hibernators, which subsist on ing of the fur. The crown of the head tends their stored fat leading to decreasing body to be black with white scatterings of hairs weight during hibernation of 30-33 percent around the nose in my northern specimens, (Grizzell, 1955). The body weights fluctuate but they were few and small. The venter throughout the year, increasing during the showed no significant difference in color, ei- growing season (for approximately six months) ther tone or pattern, and certainly was never and decreasing during hibernation. In south- whitish in the southern skins. There is slightly ern states woodchucks begin hibernation in more black suffused in the ventral fur to the early November, but they probably hibernate northward, but the character appears sporad- earlier in Wisconsin. Older and larger individ- ically in the south. It was impossible to draw uals hibernate earlier and emerge earlier (Griz- a subspecific boundary on the basis of color, zell, 1955). Aside from one record in March but it is reasonable to follow his perception and one woodchuck killed 20 December 1961 because there is probably a wide zone of in- in Portage County, Wisconsin, specimens were tergradation between the two races in Wis- collected April through September. consin. Size differences do not define the rac- The woodchuck, with long incisors and es. I follow Jackson (1961) in the following thick fur, can often protect itself from preda- classification. tors. They have been observed to chase dogs away. They snap their teeth rapidly, emit anal scent when angry or afraid, and do seem to Marmota monax monax (Linnaeus) express those emotions. Woodchucks are more intelligent than some would suspect, 1758. [Mus] monax Linnaeus. Systema naturae, ed. considering their low encephalization ratio (log 10. Type locality, Maryland. brain to log body weight). Judging from the 1780. Arctomys monax: Schreber. Die Saugethiere..., behavior of one young-of-the-year, a pam- pl. 208. Type locality, Maryland. pered house pet belonging to my wife and 1904. [Marmota] monax: Trouessart. Catalogus mam- me (Long and Long, 1965), we give the wood- malium... Suppl., p. 344. chuck credit. This young male was affectionate, playful, and on occasion tried to imitate The nominate race of the woodchuck has our behaviour even more so in some respects been ascribed to adjacent Illinois southward than a smart dog will do. After observing hu- (Hoffmeister, 1989). Inasmuch as the race is man use of the toilet seat, without any coax- characterized by larger size, I include here ing, it sat with its posterior hanging in the some measurements listed by Hoffmeister. All waste can adjacent to the stool, where he specimens were females from central Illinois defecated and urinated, then climbed down and SW Ohio. Total length, 600.3 ±49.7 and scurried away. (540-660); tail length, 134.4 ±21.7 (93-165);

TAXONOMIC ACCOUNTS / ORDER RODENTIA 163 hind foot length, 86.3 ±4.3 (78-90); greatest Waupaca Co.: Hwy 110, E Waupaca 1. length skull 93.4 ± 2.8 (90.3-97.4); zygomatic Wood Co.: Pittsville, Twn Richfield, Sect. 32, breadth, 63.1 ±3.3 (59.2-68.0); least inter- 1. Central Wisconsin 1 (not plotted). orbital constriction 25.4 ± 2.0 (23.2-28.7); nasal length 38.5 + 2.2 (35.6-41.1), maxillary tooth-row 22.1 ± 0.82 (20.6-23.1). Genus Spermophilus F. Cuvier Specimens examined. Total, 3. Dane Ground Squirrels Co.: Black Earth, 1. Dodge Co.: Taylor Road, 1 mi.S Pond Road, 1. Waukesha Co.: Douse- The ground squirrels are probably best arranged man’s Ditch, 1. following the marmots, because the thumb is primitive, usually clawed. The forefoot differs in that the thumb is functional, the index finger Marmota monax rufescens A.H. Howell. longer than the fourth, the fifth is small, and digit III is longest of all (Long and Captain, 1974). 1914. Marmota monax rufescens A. H. Howell. Black (1963) also arranged the ground squir- Proc. Biol. Soc. Washington, 27:13. Type from rels as a separate and primitive group. Elk River, Sherbume Co., Minnesota. The name Spermophilus and the common name “spermophile” were established in Measurements comparable to those giv- mammalogy until A. H. Howell synonymized en for M. m. monax for M. m. rufescens the name, preferring the older name Citellus from several northern counties are as follows, Oken. With few exceptions Oken’s names are Total Length (N = 5), 525.6 ± 44.5 (471- invalid, i.e., non-Linnaean. After a period when 578); length of tail, 4, 152.3 ± 47(120-221); both names were popular, the American Soci- hind foot length 5, 69.8 ± 10.5 (53-80); great- ety of Mammalogists followed its committee est length of skull 7, 83.5 ± 2.84 (80.4 -89.2); on nomenclature and the International Com- zygomatic breadth 8, 54.9 ±2.27 (51.5- mission on Zoological Nomenclature and con- 58.0); least interorbital breadth 8, 21.6 ± sidered Citellus invalid (Carter et al., 1968). 1.69 (19.7-24.0); length of nasals 7, 34.3 ± 1.50 (32.1-36.3); maxillary tooth-row 8, 20.5 ± 1.01 (19.4-22.0). Spermophilus tridecemlineatus (Mitchill) Remark. One adult from central Wiscon- 13-lined Ground Squirrel sin (No. 7144) has milk premolars (dP3/ ) retained. Specimens examined. Total, 18. 1821. Sciurus tridecem-lineatus Mitchill. Med. Re- Buffalo Co.: 0.5 mi. SW Mondovi 1. pos. (n.s.), 6(21): 248. Type from central Min- Calumet Co.: 3 mi. W Kiel 1. Juneau nesota (see J. A. Allen, Bull. Amer. Mus. Nat. Co.: 8 mi. N Necedah 1. Kewaunee Co.: Hist., 7: 338, 1895). T23N, R24E, sect. 20, 1. Manitowoc Co.: 1849. Spermophilus tridecemlineatus: Audubon Sect. 20, T18N, R21E 1. 1 mi. N St. Na- and Bachman. The viviparous Quadrupeds of zianz, on Hwy 131, 1. Marathon Co.: 7 mi. North America. 1: 294. from Marathon City 1. Oconto Co.: Jct. Hwys 1908. Citellus tridecemlineatus: Jackson. A prelim- T and 64, 1. Oneida Co.: Rhinelander 1.— inary list of Wisconsin mammals. Bull. Nat. Hist. 22 March, early record. Polk Co.: Clear Lake Soc., 6(1-2): 13-14. Also, 1961. Mammals of 1. U. Mn. Portage Co.: 12 mi. E Stevens Wisconsin, p.130. Point 1. Long’s home on McDill Pond (no specimen preserved). No specific locality 2. The generic name Spermophilus means Rusk Co.: Hawkins 1. Trempealeau Co.: 2 a lover of seeds. The tridecemlineatus means mi. SE Galesville, Sect. 10, T18N, R10W, 1. 13-lined, which is an appropriate name for

164 THE WILD MAMMALS OF WISCONSIN is broken, not continuous. The braincase is quite arched, especially in comparison to S. franklinii. Therefore, the foramen magnum is much more ventral in position. The baculum is ctenoid distally (Burt, 1960). There are 10 mammae on the female. Approximately 13 longitudinal stripes alternate dark brown and light tan or buff. Within and set off by the dark stripes are the Spermophilus tridecemlineatus. Phillip Grossenheider. longitudinal rows of almost square light tan spots. Either anteriorly or posteriorly some this striped and spotted squirrel. The thirteen- of these spots and rows become confluent. lined ground squirrel is known locally in Wis- The lateral dark stripes contain rows of spots consin as a gopher or striped gopher, and not as regular and definite in pattern and even has been called “picket pin”. Mitchill, an ear- the light stripes may be broken up into spots ly mammalogist working in Wisconsin and so as to resemble the other rows of spots. Minnesota, gave this squirrel its name and Approaching the belly, the lower sides are suggested it be called the Federal Ground nearly concolor with the tan or rusty and taw- Squirrel because of thirteen stripes and the ny venter. The feet are light tan or buff, with thirteen American colonies. nearly black claws. The underside of the tail Description. The thirteen-lined ground may tend toward ochraceous but the tail is squirrel is a small squirrel with a medium- bordered with brownish and distally buffy color length tail, large eyes but small ears, and a due to the buff tips and subterminal banding conspicuous pattern of stripes and dotted lines of dark brown. The upper side of the tail is that run lengthwise down the back and sides. blackish, rusty or dark ochraceous, with buff The hand or manus has five toes, and the intermixed. The flanks are often buffy or rusty thumb is short with a distinct claw. The mid- ochraceous brown. The large eyes are nearly dle or third digit is longest of the five. The black, and the postauricular area is often gray- skull is narrow, lightly built, and the molar ish or pale tan. The eyelids are conspicuously rows converge somewhat posteriorly. The pale. The forehead is irregularly and finely molars are rather low cusped, not high cusped spotted but toward the nose the fur becomes as in S. franklinii. The metaloph of the P4/ light tan. In some specimens the cheeks are pale buff, and in others the throat and cheeks are orange tawny. Some venters are pale buff whereas others are a lovely golden brown. The dark stripes vary from black to rich chocolate or dark brown, and the pale stripes vary from tan to nearly white. There are two annual molts, the summer coat (attained by June) is more brown and less grayish than the winter (attained in September). Albinistic individuals have been reported by Hoffmeister and Hensley (1949), but the spotted pattern was evident on the whitish dorsum. Kent D. Hall kept an albi- Skull of Spermophilus tridecemlineatus. Dorsal and no, also with pattern discernible, born in cap- ventral views. tivity from a mother from central Wisconsin,

TAXONOMIC ACCOUNTS / ORDER RODENTIA 165 from 1972-1976. Melanistic individuals have been noted, but none is in the Wisconsin university collections. Males may be a little larger than females. Occasionally a specimen may weigh a little over 200 g. See Table Rod-2. Dental formula. I 1/1, C 0/0, P 2/1, M 3/3 = 22. Geographic Range. The distribution dispersion of this ground squirrel has closely followed that of the fox squirrel (Sciurus niger) discussed beyond. Before the turn of the 20th century the 13-lined ground squirrel inhabit-

Maps of distribution of Spermophilus tridecemlinatus in Wisconsin and North America.

166 THE WILD MAMMALS OF WISCONSIN ed the short-grass prairies of southern and emergence, on melted ice cream or milk-egg western Wisconsin, extending northward as mixtures. A lover of sunshine, the beautifully far as Pepin, Portage, and Brown counties. patterned 13-lined ground squirrel is a won- By 1961, the species expanded its range derful neighbor scampering about the yard. northward through Wisconsin into Upper This species is widely used as a lab animal Michigan, but was unknown in Forest, Vilas, for hibernation studies. Kent Hall, emeritus and Florence counties. It was unknown north professor of this University, usually kept many of the canal at Sturgeon Bay on the Door in animal care laboratories for his studies. Peninsula. Burt (1948) recorded the species Philemus R. Hoy, about 1888, in a let- in the extreme southeastern part of the Up- ter quoted by Jackson (1961: 137) mentioned per Peninsula, and Baker (1983) had records “beneficial” habits of the thirteen-lined ground for Dickinson, Marquette, Iron, and Menom- squirrel feeding on both meadow voles and inee counties on the Upper Peninsula. The insects, over a period of 30 years.” Fitzpatrick species now occurs above Sturgeon Bay on (1925) concluded this short-grass prairie squir- the Door Peninsula. Evans (1951) suggested rel is beneficial because of the many insects that Wisconsin ground squirrels extended their consumed. range northward in the hedges and thickets Habitat. As mentioned above, the 13- associated with oldfields. lined ground squirrel prefers short-grass prai- Status. This 13-lined ground squirrel rie, but may be found in other weedy or grassy thrives throughout most of its range because communities. Most of the day it forages for Americans create short-grass fields and lawns. food. Mown areas, such as cemeteries, are Golf courses, cemeteries, mown lawns and popular habitats and provide easy hunting for parks, roadsides, campuses, and all such plac- insects. The grass may be exceptionally short, es are prime habitat for this species, even or the soil may be entirely bare. Soils are usu- though humankind wages war against them ally sandy or sandy loam, common sites in relentlessly. Their excavations are not appre- the prairies and savannas of Wisconsin. The ciated by those who like lawns and mown ground must be well drained for dry burrows. grassland. These squirrels frequently foray into On my property in central Wisconsin they the garden to eat vegetables and strawber- range into the hedgerow to co-exist with the ries, occasionally they eat bulbs and dig in eastern chipmunks (Table Rod-1), and also the flower beds, and that is about the extent into dense grassy areas while dispersing. Open of their damage. In my wife’s gardens they ground with wide visibility is as important as do less damage than chipmunks, and being food sources in the ecology of this species. rarer they are not banned from the premises. Burrows of the 13-lined ground squirrel Not all people are so tolerant. I have kept are often small outliers, but the main burrow them as pets, raised from the time of their has numerous branchings. Kent Hall (personal

Two burrows and an outlier of the 13-lined ground squirrel. After Rongstad, 1965.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 167 comm.) often found male-female pairs togeth- first litter is lost or the female fails to con- er in the burrows during the breeding sea- ceive. Perhaps that explains one juvenile spec- son. This squirrel may have multiple entranc- imen (UWSP 3718, external measurements es to its burrow. At night some ground squir- 177-64-31-6) taken 4 August in Wood Coun- rels plug their burrow entrances with soil or ty, and a remarkable record (UWSP 2995, grass. The nest is usually an accumulation of measuring 178-53-30-6) from Taylor Coun- grass. Rongstad (1965) studied and discussed ty taken on 25 September 1971. Late litters burrow systems in Wisconsin. are probably unsuccessful through severe win- Foods. The 13-lined ground squirrel ters of Wisconsin. feeds on vegetal foods such as peas, beans, Long ago as a boy in springtime, in Craw- berries, and underground tubers or roots (such ford County, Kansas, I recall that young “13- as those of beets and dandelion), but up to lined spermophiles” emerged from their bur- 70 % of the diet may be insects (Streubel and rows in June. In Wisconsin, at least in central Fitzgerald, 1978). Many kinds of seeds are Wisconsin, they emerge in early July. In south- eaten, often stuffed into the mouth, which ern Wisconsin emergence of young was ob- opens into paired internal cheek pouches, for served 20-23 June, and the young begin to transport of food to the burrow. Some plants disperse eight days later. One (UWSP 1910) used in this way include sunflower, ragweed, was taken near Stevens Point 12 June. Kul- dandelion, vetch, grasses, and knotweed. The pa and Hall (1983) isolated 28 pregnant fe- animal foods include insects, such as grass- males from Wisconsin Rapids, and they gave hoppers, crickets, ants, beetles and their birth from 25 May to 1 June. grubs, stoneflies, and true bugs. Worms and Gestation lasts from 27 to 28 days, and millipedes also are eaten. Opportunistically, the young are born naked, blind and helpless the young of birds (as well as eggs of ground (Streubel and Fitzpatrick, 1978). Usually there nesters), small lizards or snakes, and even are about 8 offspring, but there can be as many small mammals are eaten. In late June (2002), as 13 (Zimny, 1965). Rongstad (1965) found Joseph Graceffa, in Stevens Point, Wiscon- 8.7 ± 0.03 litter size in Wisconsin. Newborn sin, observed a 13-lined ground squirrel leap young weigh only 6-7 grams. By 12 days the from the ground to capture by the neck a pattern of spots and stripes is visible on the young (feathered-out) house sparrow (Passer now darkened skin and the hair is beginning domesticus) as it attempted to leap into flight. to grow out. Soft trilling calls are made. By 20 It ate the brain at the site of capture and left days the incisors emerge and the young crawl the bird’s carcass on the ground. This ground about. By 26 days the eyes open. Weaning squirrel also feeds on carrion, even carcasses was complete after 5 to 7 weeks. of its own kind (Bailey, 1893; Howell, 1936; Only a third-grown, the young venture Whitaker, 1972a; Yeager, 1937; B. Bailey, from their burrows and search for plant and 1923; Brigwater and Penney, 1966). possibly insect foods. They soon dig their own Reproduction. Males are reproductively burrows, small and shallow at first. By the time active when they emerge from winter hiber- for hibernation, although they delay it a few nation. Females are viable a few days later. weeks, the young are only 3/4 grown. Bridg- Ovulation seems induced (Streubel and Fitz- water (1966) discussed growth and develop- patrick, 1978). Multiple paternity has been ment in this species. Longevity is approxi- noted (Hanken and Sherman, 1981). Partu- mately 7-8 years (Crandall, 1964). rition begins in mid-April and continues until Mortality. Aside from humans, who trap early May in Wisconsin; there is a single litter and poison this species in lawns and fields, per year (Rongstad, 1965). There are a few and kill thousands with automobiles along the cases of second matings, perhaps when the highways, the badger is an important preda-

168 THE WILD MAMMALS OF WISCONSIN tor (Leedy, 1947; Long and Killingley, 1983). es. In late September the fat adults (especially Weasels, domestic cats, dogs, foxes and coy- adult males) begin to stay underground, and otes, and various hawks eat them. Even those not so fat may continue feeding anoth- shrikes crows, and prairie snakes eat them er month before entering hibernation. Kulpa on occasion (Craighead and Craighead, 1956; and Hall (1983) reported that males attain Linduska, 1950; Dearborn, 1932; Jackson, greater body weights than females but enter 1961; Fitzpatrick, 1925; Schofield, 1960). hibernation about the same time. Streeter and Parasites include blood protozoans, cestodes, Hall (1983) found in weaned squirrels that nematodes, spiny-headed worms, botfly lar- various diets led to varying peak weights, but vae, mites, ticks, lice and fleas (Jackson, 1961; for each of these a hypophagic period of di- Scharf and Stewart, 1980; Whitaker, 1972; minished body weight followed, leading to Streubel and Fitzpatrick, 1978). hibernation. In young-of-the-year the onset Home Range and Density. Gunderson of hibernation was in late September, even in (1976) in Minnesota reports home range for juveniles treated with the appetite depressant males up to 11.7 acres, and 3.4 acres for naloxone. However, I observed one foraging females. Distributions of 13-lined ground for food as late as 16 October 2002. Sex, squirrels are patchy, because short-grass prai- diet, and maximum weight seemed to have rie is patchy. McCarley (1966) found one rea- no trigger effect on hibernation, but after peak son autumn populations are limited is that weights were attained followed by the hypoph- young of the year disperse in late summer. agic phase, hibernation soon followed. Curled Adults ranged from one per three to one per into a ball, in a nest of grasses, the squirrel 4.5 acres. After weaned young appeared falls into a deep dormancy and its metabo- above ground, the number increased to 2 per lism falls. Such winter sleep is defined as hi- acre. Evans (1951) found in one summer 30- bernation when, and only when, the temper- 35 ground squirrels in a 15-acre field. In Ohio, ature fluctuates markedly with the ambient Lishak (1977) observed as many as 12 adults temperature. In S. tridecemlineatus the body and juveniles per acre. There are some favor- temperature may drop, so long as the envi- able habitats, such as in cemeteries and golf ronmental temperature drops, to about 3o C, courses, where Wisconsin 13-lined gound which is a few degrees above freezing of wa- squirrels are more abundant than 2 per acre. ter. The heart rate drops from 50-350 beats Jackson (1961) suggests as many as 20 per per minute to a bradycardia of only 5 per acre in some habitats. Rongstad (1965) found minute. Breathing rates drop from 50 to 4 annual mortality high, 81 percent. times per minute. Because the metabolism is Remarks on Hibernation. Although the so much reduced, the hibernation fat is con- 13-lined ground squirrel is not the only hiber- sumed very slowly. The squirrel remains alert nator in Wisconsin, it was studied earliest and to physical stimuli such as touching or stretch- has been studied most intensively. P. R. Hoy ing, but otherwise it seems asleep. There (1875) discussed it first. G. E. Johnson and seems to be only one ground squirrel per hi- his students made many early studies, and bernaculum burrow. Johnson and Hanawalt (1930) wrote a time- Sometimes the ground squirrels arouse less classic on the subject. Other early papers from deep sleep, even to awake and emerge were by Wade (1927, 1930). A more recent above ground in southern latitudes. This is work is by Fisher and Mrosovsku (1970). This unknown in Wisconsin, where winters are species exhibits a circannual rhythm involv- severe. Beer (1962) reported that they emerge ing hibernation, and also fluctuations of me- in Minnesota as early as 26 March. Males tabolism, reproduction (in spring), weight gain emerge before females, and are immediately (in summer), and other physiological chang- ready for courtship and breeding. They were

TAXONOMIC ACCOUNTS / ORDER RODENTIA 169 prepared for reproduction even while in their months). The metabolism is so low in the burrows, but the females do not enter estrus Wisconsin squirrels it seems doubtful that they until emergence. Wade (1950) suggests that eat, especially without water available. the ground squirrels cannot emerge until the Geographic variation. Because the race ground thaws, but since chipmunks of simi- is the nominate race, probably little attention lar size and form may emerge through black- has been paid to its geographic variation. There top roads and sometimes clamber about on is a great deal of microgeographic variation deep snow in Wisconsin, I must question this from place to place in pallor and venter colors suggestion, especially in southern parts of in Wisconsin, but no significant speciation. Wisconsin and in sandy soils of central Wis- Probably many of the northern populations consin. Obviously the return to normal me- emigrated from southern ones, so it seems tabolism requires a great expenditure of en- unlikely that in recent times any differentiation ergy, which relates to considerable weight would have occurred. Some populations are loss, sometimes almost half the weight in quite pale, whereas others are dark brown. August. The body weights are lowest, in Some individuals show reddish brown and the adults, during spring reproduction (following venter may be variably buff or golden tan. soon after hibernation). Specimens examined. Total, 80. Ad- Hibernation seems a great strategy for a ams, Bayfield, Columbia, Dane, Dodge, mammal of warm habitats to invade the north- Door, Douglas, Dunn, Green, Iron, Jeffer- ern regions, such as Wisconsin prairies (which son, Juneau, La Crosse, Lincoln, Manito- are frost bound to depths of down to five feet woc, Marathon, Marinette, Monroe, Ocon- in winters of little snow). It is therefore odd to, Oneida, Pepin, Portage, Price, St. that this species hoards stores of seeds but Croix,Taylor, Walworth, Waukesha, hibernates so much of the year (almost six Waushara, Wood counties.

Table Rod-2. Some external and cranial measurements of adult Spermophilus in Wisconsin.

Spermophilus tridecemlineatus S. franklinii Sex and N M, 8 5 3 2 F, 6 4 1 – F, 4 Total length 247.4±19 264.8±18 284 245, 286 – 246.5±16 241.3±32 286 – 360.3±37 Tail length 83.3±10.6 95.4±4.3 93 65, 74 – 89.5±10 87.5±7 98 – 124.7±31.2 Hind foot 34.3±1.8 36.4±1.3 38.7 34, 33 – 32.0±2.8 36.0±1.6 39 – 51.5±2.4 Ear length 6.83±1.4 10.8±2.2 6 11, 9 – 6.6±2.1 9.75±0.5 5 – 12.0±1.15 Greatest length of skull 40.1* 40.75±2.3 40.5 38.6, 39.2 – – 40.73±0.95 42.5 – 52.8±0.15 Zygomatic br. 23.9* 22.6±1.8 23.4 22.9, 21.6 – – 22.7±0.6 24.6 – 29.5±0.48 Interorb. Br. 7.6* 7.4±0.5 7.9 7.2, 7.4 – – 7.58±0.4 8.3 – 10.44±0.39 Length nasals 12.4* 12.67±0.97 13.2 12.7, 10.3 – – 12.1±0.8 13.9 – 18.75±0.14 Maxillary toothrow 7.13* 7.56±0.5 7.5 7.0, 7.0 – – 7.89±0.4 7.4 – 9.9±0.28 *N = 4

170 THE WILD MAMMALS OF WISCONSIN Spermophilus franklinii (Sabine) Franklin’s ground squirrel is larger, long- Franklin’s Ground Squirrel er-bodied and much longer-tailed than the 13- lined ground squirrel S. tridecemlineatus. The 1822. Arctomys franklinii Sabine. Trans. Linnaean tail is bushy and almost half the body length. Soc. London, 13:537. Type from Carlton The skull is long and narrow, especially the House, Saskatchewan. nasals and rostrum. The braincase is more flat- 1827. Spermophilus Franklinii: Lesson. Manuel de tened than in S. tridecemlineatus, and the Mammalogie ... p. 244. width between the orbits is wider. The teeth 1938. Citellus franklinii: A.H. Howell. N. Amer. (especially the lower molars) have higher cusps Fauna 56:57. and cuspids than in S. tridecemlineatus, probably for eating flesh. The ears are low and The great anatomist F. Cuvier in 1825, rounded, unlike that of the gray squirrel (which not his famous brother Georges Cuvier, es- has a similar whitish but even bushier long tail). tablished the generic name Spermophilus. The claws of the forefeet are short and robust. This word means seed-lover. The naturalist There is a claw on the pollex rather than a nail Sabine named this squirrel in honor of the or “hoof,” seen in tree squirrels, but five toes famous British explorer of the Arctic, Sir are on each foot. There are large internal cheek John Franklin, who was lost in the Canadi- pouches, 10 mammae, and the chromosome an Arctic in 1846. See status of Oken’s name pattern is 2N = 42 (Nadler, 1966). Citellus in account of Spermophilus tride- The skull of Spermophilus franklinii is cemlineatus. much larger than the skull of the 13-lined Description. In the western states there ground squirrel, but similar. Dorsally, the pos- are many diverse ground squirrels that share torbital processes flare away from the cranium with this species a typical coat pattern of light and are quite long. The jugal extends notice- brown, tan or buff upperparts dappled with ably behind the zygoma, more so than in the small spots of whitish or buff. Such a pattern 13-lined ground squirrel, and the lambdoid is cryptic coloration in grassy habitats, where crest is higher in mature animals. The ventral sunshine is filtered by the grasses or sage, and aspects of the skull are very similar, except the the ground is dappled with sunlight. This spe- teeth, which are more massive in S. frankli- cies is the only ground squirrel in Wisconsin nii. From the side, the skull is more flattened, exhibiting such a pattern, and the flecking is not nearly so arched as in the 13-lined ground never mentioned in most descriptions. squirrel. The rostrum is relatively longer and

Drawing of Spermophilus franklinii. V. Hogg, Univ. Skull of Spermophilus franklinii. Dorsal and ventral Kansas (in E. R. Hall). views.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 171 the zygomata angled posteriorly more acutely song birds and duck nests, this species is harm- than in Sciurus, resulting in a narrower, trian- less insofar as crops and gardens are con- gular anterior plate for origin of the masseter, cerned. Since it avoids mown, grazed, and instead of a nearly four-sided plate in Sciurus. well-drained pastures it does not come into When the skull rests on its dorsal sur- contact much with humanity. It is seldom face, the foramen magnum opens upward in abundant, but occasionally is found in loose Sciurus but opens posteriorly in S. frankli- breeding colonies of four to eight individuals. nii. The baculum is toothed distally, as in the At the margin of its range, in Central Wis- 13-lined ground squirrel. consin, mostly in Portage and Wood coun- The upper parts are brownish gray in- ties, one or two of these squirrels were taken termixed with a suffusion of yellowish and every one or two years from 1967-1972. black, dappled lightly with tiny specks of din- None has been collected since 1972. Recently gy white. The eyelids, cheeks, throat, and in- Lewis and Rongstad (1992) reviewed the sta- ner legs are buffy. The tail is whitish inter- tus of this squirrel in Illinois and Wisconsin. mixed with buffy gray but with a pale creamy Habitats. Franklin’s ground squirrel prefers suffusion visible. The head and adjacent neck marshes and tall-grass prairie, especially along- is a pronounced silvery gray, the hairs lightly side wooded areas, railroad rights-of-way, and tipped with white. There is a single molt in weedy fields. Mowing prairie is adverse to this early summer, and winter pelage is bleached. species, which is found in tall weeds, grasses Observed weights range from 450-460 and forbs, including clover, dock, big bluestem, g. Measurements are given in Table Rod-2. other grasses, thistles, goldenrod, dandelion, Dental Formula. I 1/1, C 0/0, P 2/1, M strawberry, cocklebur, wheat, and so forth. 3/3 = 22. Standing water is often nearby. The species has Geographic Range. Franklin’s ground been taken near sand dunes, especially on rail- squirrel inhabits marshes and wet meadows road embankments, where it often makes its in the western and southern prairie counties, burrow system, and on weedy roadsides (Mum- but is not known in the extreme southwest- ford, 1969; Jackson, 1961; and others). ern corner, nor in most counties along Lake The burrow system has been described Michigan. It ranges as far northwest as Polk by Kennicott (1858 or 1859) and Ellis (1982), County and in the center to Clark and Por- as longer and deeper than that of the 13-lined tage counties. The species probably occurs in ground squirrel, often sited on a bank in weedy Marathon County (Mead Wildlife Area). cover, and often having the entrances plugged Status. Never abundant, this unusual with loose dirt. The diameter of the tunnels squirrel is on the Special Concern list of Wis- averaged 8-10 cm. consin species, and might be diminishing in Foods. A variety of grasses, herbs, ber- numbers. Sowls (1948) reported that this ries, and seeds is eaten. Plants include sow ground squirrel accounted for 19 % destruc- thistle, chokecherries, elderberries, beach pea, tion of duck nests studied in Manitoba. In cen- and clover (Sowls, 1948; Seton, 1953; Long, tral Wisconsin I know that no such predation 1967). About 1/3 of the diet is animal mat- on this scale exists. Based on my observa- ter. Numerous insects such as beetles, grubs, tions, the snapping turtle (Chelydra serpen- caterpillars, grasshoppers, crickets, ants and tina), common crow (Corvus brachyrhyn- their eggs are eaten (Bailey, 1893). Seton chos), and mink would kill many more duck- (1953) noted feeding on flesh: deer mice, lings and eat more eggs robbed from the nests young cottontails, and chickens. I have ob- than S. franklinii. This ground squirrel is served them in Kansas feeding on young dick- never abundant in Wisconsin. Although not cissels and their eggs, and once in Illinois (near entirely beneficial because of its predation on Urbana) on a road-killed pheasant (Long,

172 THE WILD MAMMALS OF WISCONSIN 1986b). Sowls (1948) discussed their predation on duck eggs and ducklings in Manitoba. Reproduction. Gestation is about 26-28 days, following April courtship, which takes place as the squirrels emerge from hibernation. The young are born in May or June, and lactation commences then continuing until weaning some 40 days later. The single litter size varies, from 5-11 young. The young are born naked and blind; hairs appear on their bodies in 10 days. The eyes open in 20 days. They begin to “whistle” in 20 days and emerge from the burrow in approximately 30 days. Weaning is a few days thereaf-

Maps showing geographic distribution of Spermophilus franklinii in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 173 Sketch of Eutamias minimus by E. T. Seton. ter. One Franklin’s ground squirrel lived 7 years ter breeding they are seldom seen, apparent- and two months (Crandall, 1964). ly beginning hibernation as early as late Sep- Mortality. The Red-tailed Hawk is the tember. Dates on my specimens, indicating primary enemy, according to Seton (1953). summer activity, range from 12 May, with Foxes, coyotes, striped skunks, mink, wea- lactation on 15 July, two specimens in late sels, and an occasional hawk or owl catches September, and one as late as 15 October. them (Baker, 1983). The badger, of course, Hibernation, therefore, may not last seven is a specialist in ground squirrel predation months as reported by Hoffmeister (1989) in (Long and Killingley, 1983). Large snakes may Illinois. Hibernation reportedly lasts from Sep- eat them. Jackson (1961) mentions lice, fleas, tember to April, the males appearing above protozoans, tapeworms and roundworms. ground a week before the females. Many are killed by cars. Insecticides and weed Geographic Variation. None is evident killers may have caused the species to decline. in Wisconsin. Home Range and Density. There is noth- Specimens Examined. Total 8. Dane Co. ing reported to my knowledge on home range. Grove W. Area 1 UW Wild. Ecol. Portage The Franklin’s ground squirrel is colonial or Co. Sect. 2, T23N, R7E, 2. 1 mi. W Hwy; N, at least forms “loose communities” for rear- 1. 1/2 mi. W Junction City, 1. 2 mi. S Junc- ing the young (Hall, 1955; Gunderson and tion City, 1. Jordan Swamp, 1. Wood Co. 3/ Beer, 1953). Populations peak perhaps ev- 4 mi. SE Arpin 1. ery four to six years with as many as 30 animals per acre (Schwartz and Schwartz, 1981). Sowls reported 14 shot in less than an hour Genus Eutamias Trouessart at a locality in Manitoba. Banfield suggests 4- Western Chipmunks 8 per acre (1974) is common. Remarks on Habits. Diurnal, secretive, I have studied the arguments presented for except when calling back and forth to one arranging all the chipmunks of North America another or chasing one another in breeding, in the genus Tamias Illiger and am uncon- the Franklin’s Ground Squirrel has a low, vinced. Hoffmeister (1989) and Baker (1983) musical trill which signals its presence. It hi- also choose to retain the name Eutamias for bernates in winter, and spends much of the the western chipmunks. I am most persuaded summer underground as well. Prior to hiber- by the argument of John White (1953). Taken nation fat is deposited on the body, which together with his other works at that time it lasts the animal until it emerges in April. Af- comprises a scholarly and intelligent classifica-

174 THE WILD MAMMALS OF WISCONSIN tion of chipmunks in relation to the other Sci- least, referring to small body size. This squirrel uridae. White carefully weighed the usefulness is the smallest of Wisconsin’s Sciuridae. of characters, which in squirrels are notorious- There is a single subspecies in Wiscon- ly polyphyletic and convergent in their evolu- sin, which Jackson referred to Eutamias min- tion, choosing those in which he held the stron- imus jacksoni (A.H. Howell) without com- gest confidence. Some of these had been dis- ment. Other authors followed suit. The name paraged owing to “non-functional” persistence that should be applied to this race is Eutamias (upper third premolars) but he found them func- minimus neglectus (J.A. Allen). Allen exam- tional (showing tooth wear). Loss of the P3/ ined specimens from Upper Michigan (near in Tamias may have evolved in response to Escanaba), northern Minnesota, and the root competition of the adjacent premolars. northeast shore of Lake Superior referring That process has adaptive significance. It is them to a bright colored race from the Lake important that Tamias has as its type species Superior region. The last locality mentioned the eastern Tamias striatus, so that the tax- became the type locality (see below). Subse- onomy of the North American Eutamias dif- quently, Howell drew a line between the bright fers geographically from that of Tamias. White chipmunks of the Lake Superior region and a listed ten characters, including for Tamias the northern race Eutamias minimus borealis, elongated head of the malleus, sharp angle so that the type locality of neglectus fell in between lamina and manubrium of malleus, the range he ascribed to borealis. Therefore, ventral keel on tip of baculum (resembling Sper- each worker had a different opinion on the mophilus), unfused hypohyal and ceratohyal identity of the chipmunks of the northeast bones, rounded tendon of digastric muscles, shore. Howell assigned all the chipmunks absent upper P3/, lingual anterior root of P4/ examined from the north shore to the bright , short tail (as in Spermophilus), medial white race. Since he excluded the type locality of stripes twice as wide as the others, lateral pairs neglectus from the range thus delineated for of white stripes all short. Some of these char- the bright chipmunks, neglectus became a acters are consequential of one another. They synonym of borealis. He then renamed the persuade me to retain both the name Eutamias bright chipmunks jacksoni. The concept of and the name Tamias. Recently, Ellis and borealis at that time was broader than now, Maxson (1979) obtained immunological evi- for example it extended into Wyoming. Ander- dence to support this thesis. Burt (1960) said son (1946) drew the line between the bright there is no evidence from bacular form that chipmunks and borealis much farther north- Eutamias and Tamias are one genus. ward, and that explains better the presence of bright chipmunks all through the woodlands north of Lake Superior (even as Howell ad- Eutamias minimus (Bachman) mitted), which some of my specimens con- Least Chipmunk firm (see below). Therefore, the name jacksoni is a synonym of neglectus. 1839. Tamias minimus Bachman. J. Acad. Nat. Sci. Philadelphia, 8:71. Eutamias minimus neglectus (J.A. Allen) The Chipmunk. He moves with flickering indecision Like strange stripes across the television. — 1890. Eutamias quadrivittatus neglectus J.A. Allen. Ogden Nash. Bull. Amer. Mus. Nat. Hist., 3:106. Type from Mouth of the Montreal River, Ontario. The name Eutamias means a true hoarder of 1922. Eutamias minimus neglectus A.H. Howell. stores (or foods), and minimus of course means J. Mamm., 3:184.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 175 1925. Eutamias minimus jacksoni A.H. Howell. J. The total length is about 200 mm. The Mamm., 6:53. Type from Crescent Lake, Onei- tail is relatively longer than in Tamias, 40 da Co., Wisconsin. percent or more of total length. Weights vary 1944. Eutamias minimus neglectus: Anderson and up to 50 g. See Table Rod-3. Rand, Candian Field-Nat., 57:134. Also Ander- Dental Formula. Note tiny upper third son, Nat. Mus. Canada Bull., 102:114, 1946. premolars; the formula is I 1/1, C 0/0, P 2/ 1, M 3/3 = 22. Description. The skull generally resem- Geographic Range. The least chipmunk bles that of Tamias, narrow but broad across is found in suitable habitats in forested north- the interorbital constriction. The presence of ern counties, extending southward as far as a tiny upper third premolar on each side dis- Juneau, with recent occurrence in Sheboyg- tinguishes this tiny squirrel from Tamias. an County. Bright stripes, small body size and a relatively Status. Jackson (1961) considered the long tail also identify this quick and bright- least chipmunk as abundant as the eastern eyed chipmunk. The median dorsal black chipmunk. He was wrong, I have looked in stripe is the longest extending from the brown- the wrong habitats, or Eutamias has dramat- ish nape or crown along the back onto the ically declined. The species seems to have base of the tail. Adjacent to the median stripe vanished in some of the southern parts of its is a narrow strip of pelage brownish or gray- range, possibly in competition with Tamias ish and this fur is bordered laterally by a bright striatus, even though Reilly (1970) found no white (occasionally buffy or creamy) stripe competition between these species on the which is enclosed above and below by con- nearby Upper Peninsula of Michigan. Some spicuous black stripes. Lateral to these on the suspect competition between the two chip- sides of the chipmunk is beautiful ochraceous munks because Tamias becomes so abundant or cinnamon buff pelage, which is bordered in proximity to man’s habitations. Another below by the pure white venter. The dorsal competitor might be the red squirrel. stripes, even the lateral ones, are elongate, On the Upper Peninsula clear cutting in extending to the base of the tail. A rusty chest- the forests seems to be benefiting the least nut or brown patch at the base of the elon- chipmunk, according to Verme and Ozoga gate tail present in Tamias is lacking in Eu- (1981). The animal is a decided asset for the tamias. Chipmunks have stripes on the head, Wisconsin fauna, being one of the most col- and often in Eutamias minimus the stripes orful, cutest, and friendliest denizens of camp- are conspicuous. The white of the eyelids sites in the North Woods. The damage in some below and above the eye extend anteriorly cabin or to some garden or crop is hardly to the nose, and posteriorly almost to the significant. In Wisconsin it has no financial ear. Post-auricular patches of gray are evi- impact. To obtain a measure of general abun- dent. The tail is grizzled grayish brown or dance one might compare the specimens pre- brownish gray above, and bright ochraceous served in this collection (about 200 Tamias brown or ochraceous below. The feet are to 48 Eutamias). similar to the brown or gray dorsal pelage, Habitat. Often reported from conifers, that strip which is not involved with striping. and rocky lakeshores, the least chipmunk is The bacular tip has a tiny dorsal keel, and most likely to be found in northern counties the form of it is slender. There are eight (two in wetlands, such as conifer swamps and ri- pectoral, four abdominal, and two inguinal) parian habitats of willows and alders. It is of- mammae. Internal cheek pouches are large ten found, with Tamias, at the forest edge, in this species. Winter pelage is more gray- where visibility is good, soil black, and natu- ish, as is the juvenile pelage. ral foods available. If there is ecological sepa-

176 THE WILD MAMMALS OF WISCONSIN ration between Eutamias and Tamias, in Ground cover would seem less significant Wisconsin or the Upper Peninsula, the least in dashes by this quick-moving chipmunk over chipmunk is found in wetter habitats. In Alg- stony habitat, except for scatterings of brack- er and Schoolcraft counties, upper Michigan, en fern, blueberries and raspberries, with Ozoga and Verme (1968) found numerous stunted jack pines often present. Some brush Eutamias in conifer swamp, and only a sin- and other cover are probably necessary for gle Tamias was taken with them. Tamias foods and security. prefers higher, dryer home sites. Both spe- The chipmunk excavates its burrow and cies occupy hardwoods, conifers, and forest the opening is often plugged, while a new main edge together, especially in rocky places. Don entrance is inconspicuous. The burrow may be Follen, in October 1967, collected least chip- worked between and among rocks. Nests may munks in oak-maple hardwoods mixed with be constructed in the burrows, or hollow trees, some aspen near Alvin in Forest County. One woodpecker cavities, and stumps, or beneath was trapped in a grassy marsh nearby. I ob- fallen logs, in rock piles, and perhaps on occa- served them on several occasions on granite sion as accumulations of grass and leaves in outcrops of lakeshores, with jack pine, blue- trees. Orr (1930) found one such summer nest berries, mosses, lichens and sparse grasses. 18 feet above ground in a black spruce. Usually there were spruce and aspen also. Winter nests are often constructed above In western North America, at various el- hoarded foods, at one observed depth about 3 evations, least chipmunks often inhabit mo- feet beneath the ground surface. These nests raines, sage covered gullys, and even flat are comprised of shredded bark, grasses, cat- desert places, as well as the montane forest kin fibers, and even mammal hair and feathers. habitats. In Wisconsin and the Upper Penin- Foods. Little is known about foods of sula there are no habitats resembling those least chipmunks. Probably they eat the same warm, dry and exposed soils, even though kind of foods the other chipmunks eat, namely Eutamias is, since the Pleistocene epoch, an seeds, grasses, fruits, insects, perhaps an oc- invader from the far west. There is rocky till casional egg (Criddle, 1943; Vaughan, 1974; on Rib Mountain and its nearby hills, resem- Forbes, 1966). Blueberry seeds (befitting wet bling talus in the mountains of the Rockies. habitats), smartweed and timothy seeds have Eutamias once occurred there, but probably been found in cheek pouches (Banfield, the little chipmunks had ranged along the Rib 1974), and a cache for winter contained hun- River (at the foot of the mountain) where the dreds of acorns and thousands of cherry pits. species still occurs upstream near Athens. Hazard (1982) reported that they ate strawberries. They often eat raspberries. These berries often grow in seral or edge habitats. Reproduction. Following their long winter sleeping, least chipmunks breed while chasing each other above ground. In early May, pregnant females construct nursery dens with nests of grass; after an estimated 31-day gestation period 5 or 6 (2-7) young are born, usually in early June. They are naked, the eyes and ears closed. Each neonate weighs about 2.3 grams. The eyes open in about 28 days, and the young are well haired by 40 days. In less than two months the young are weaned. Usually a single Skull of Eutamias minimus. litter is born; late litters probably result from

TAXONOMIC ACCOUNTS / ORDER RODENTIA 177 loss of an earlier one (Skryja, 1974). Young-of- the-year apparently do not breed. Mortality. Little is known of predation on least chipmunks, especially in Wisconsin. The usual hawks, owls, perhaps crows and ravens, possibly large fishes, snakes, and Carnivores must be important predators. Weasels and martens are suspected to kill them. Seton pictured such a “wanton” killing of a chipmunk family by a weasel. Probably many fail to emerge after a long winter, if cached food was in short supply. Forest fires doubtless destroy many, although burnt over forest land

Maps showing geographic distribution of Eutamias minimus in Wisconsin and North America.

178 THE WILD MAMMALS OF WISCONSIN features the reappearance and abundance of Bayfield, Douglas, Florence, Forest, Iron, Eutamias after a few years. Juneau, Langlade, Marathon, Oconto, Onei- Parasites include internal nematodes da, Sheboygan, Taylor, Vilas counties. (Jackson, 1961) and arthropods such as fleas, Michigan. Delta and Schoolcraft counties. ticks, mites, and lice (Jackson, 1961; Man- ville, 1949; Timm, 1975). Home Range and Density. Little is known Genus Tamias Illiger about these parameters in least chipmunks. Eastern Chipmunk Jackson (1961) speculated 30 or more per acre in suitable habitats. I have never seen The characters distinguishing Tamias from them more numerous than 5-8 in an acre, Eutamias were discussed in the account of and quite localized within that acre while ab- Eutamias. The eastern chipmunk is larger, sent in adjoining habitats. Reilly (1983) sug- less brightly colored, and the dorsal stripes gests 5-15 per acre as more usual values. do not extend to the relatively shorter tail. Reilly (1970) and Manville (1949) found from The upper P3/ is lacking, there is a chestnut less than one to 5.6 per acre on the Upper or rusty brown patch on the rump, and the Peninsula. Individuals wander over home rang- tip of the baculum has a minute ventral keel. es of less than an acre to 4.3 acres. Remarks on Behavior. Diurnal, the least chipmunk vocalizes with sharp calls for danger and growling among themselves. Usually when they call they jerk their tails. Not as territorial as Tamias, the adults nevertheless seem to maintain spatial separation (Reilly, 1970). The winter is passed in winter sleep (sometimes regarded as torpor, because they are often active above snow and underground). The animals disappear underground from Oc- tober until March. Additional Natural History. Forbes (1966) studied least chipmunks in northern Minnesota. Geographic Variation. There is no ap- Eastern Chipmunk. Sketch by anonymous student, parent geographic variation in Wisconsin and Wildlife Society Newsletter, UW-Stevens Point. the Upper Peninsula of Michigan. Further, specimens from the north shore of Lake Su- perior are no different, except that a series of five from 25-30 miles northwest of Kenora, Ontario were a little grayish. Fresh pelage observed on chipmunks from Horseshoe Lake, Manitoba, was bright. These and all other least chipmunks on the Upper Peninsula are referred to Eutamias minimus neglectus (J.A. Allen) because of their characters and on geographic grounds. See argument above. Specimens examined. Total, 48. From Upper Michigan, 4. Wisconsin. Ashland, Skull of Tamias striatus.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 179 Other traits are included below in the account proached me, jumped up and tried to snatch it.” of the Wisconsin subspecies. There is a sin- — William Brewster, quoted in E. T. Seton’s Lives gle species in this genus. Perineal, anal, and of Northern Animals, 1909; 1953, II, p. 201. oral glands were discussed by Snyder (1982). The baculum (Burt, 1960) averages about 4.15 mm in length, and there is an os clitori- Tamias striatus (Linnaeus) dis. The skeleton and musculature are described by Bryant (1945). The feet are penta- 1758. [Sciurus] striatus Linnaeus. Systema naturae, dactyl except the thumb is vestigial and bears 10th ed. 1:64. Type locality fixed by A.H. a small nail. The tail is short, about 30-38 Howell, N. Amer. Fauna, 52:14, 1929, Up- percent of the total length. There are four per Savannah River, South Carolina. pairs of mammae. 1857. Tamias striatus: Baird. 11 th Ann. Report, Smithsonian Inst., p. 55. [The Eastern chipmunk] When (1888) the 1891. Tamias striatus griseus Mearns. Bull. Amer. famous Cambridge naturalist William Brewster Mus. Nat. Hist., 3(2): 231. Type from Fort shot and broke the wing of a wood thrush and it Snelling, Minnesota. went fluttering along the ground, a Chipmunk 1932. Tamias striatus lysteri: Komarek. J. Mam- pursued and caught it. “The Chipmunk killed mal., 13: 207. Also Barger, Wisconsin Cons. the bird and had eaten most of the brains... As I Dept., Publ. 351, p. 11, 1951. held the bird dangling, the Chipmunk ap-

Table Rod-3. Standard external and cranial measurements of Wisconsin chipmunks Tamias striatus and Eutamias minimus. There are no significant differences among the populations of Tamias striatus.

Locality N Sex TL Tail Hind foot Ear Gr. l. Zygo. br. Int. br. l. nasals Max. t-r Tamias striatus Bayfield Co. 9 M 248 96.4 37.22 17.11 40.7 22.6 9.98 13.09 6.16 +12.5 +11.9 +2.1 +2.6 +1.1 +0.67 +0.36 +0.58 +0.53 2 F 235 83 34 19 39.95 23.06 10.69 12.13 6.14 248 90 36 19 41.5 22.9 11.55 12.89 6.10 Portage Co. 5 M 250 92.6 35.25 21.2 4.94 22.64 10.14 13.79 6.20 +15.0 +7.8 +1.26 +6.7 +0.39 +0.42 +0.63 +0.82 +0.21 6 F 236 86.67 33.17 16.17 40.33 22.03 9.79 12.31 5.95 Vernon Co. 3 M 268 83 33 17 42.65 23.4 10.35 13.65 5.89 245 102 30 18 230 97 31 14 Grant Co. M 265 102 36 16 41.32 23.0 10.62 13.14 6.1 Dodge Co. 5 M 242.5 87.5 33.75 15.25 41.4 23.0 9.91 12.9 5.96 +21.7 +17.0 +2.0 +1.5 3 F 221 90.7 35.3 17.3 40.47 22.0 9.54 13.05 5.94 Racine Co. M 266 84 34 12 41.9 22.8 10.2 13.4 5.6 Tamias striatus doorsiensis (after Long 1971) Door Co. 5 Adults 259 95 36 20 — 23.5 — — 6.6 8 sub-adults — — — — — 22.6 — 13.9 6.6 Eutamias minimus Forest Co. 6 M 198.0 85.4 29.0 13.8 31.29 17.79 6.82 9.41 4.9 +8.0 +7.7 +1.58 +1.6 +0.96 +0.57 +0.18 +0.86 +0.23 2 F 198 82 28 12 31.8 17.8 7.05 9.45 4.7 197 80 28 13 31.3 17.65 6.8 9.76 4.94

180 THE WILD MAMMALS OF WISCONSIN The name Tamias means a hoarder and Geographic Range. This species is one steward of stores (or foods) and striatus means of the commonest mammals in Wisconsin, striped. The names griseus and doorsiensis re- occurring in every county and in a wide range fer, respectively, to gray and to the land of Door. of habitats. It is even abundant in human set- Description. Same as set forth above for tlements. It is not found on the Apostle Is- the genus. The skull is broad across the inter- lands or on the Grand Traverse islands (in- orbital constriction in both Wisconsin races, cluding Washington, Rock and other isles in and exceeds that of chipmunks in lower Mich- Door County). It also avoids especially wet igan. The two races in Wisconsin do not dif- and prairie habitats, and is seldom seen in fer much from one another in this respect. the “goat prairie” region of western Wiscon- The chromosomes number 2N = 38, with a sin. See Map. dot-like Y chromosome (Snyder, 1982). Status. This colorful, lively little squirrel Other Wisconsin squirrels of comparably is abundant and maintains high densities in small size, including the Spermophilus, are suitable habitats. It thrives even in the prox- never striped on the head, except Eutamias, imity of humankind. It may become a pest on which chipmunk is similar. The dorsal, longi- garden fruits and vegetables, feeding also on tudinal stripes in Eutamias extend to the base flower garden bulbs, and caving in lawns. After of the tail. Tamias features a mid-dorsal stripe spending quite a sum on an asphalt driveway, of dark brown dividing the dorsal grizzled pel- three chipmunks burrowed under it and age (grayish, brownish or even ochraceous in caused deep snake-like depressions as their tone). Laterally are two narrow brown stripes burrows caved in. Twice in early spring, the enclosing a whitish or buffy white stripe. This chipmunks burrowed up through the asphalt pattern is seen on either side. The rump is road in front of our house. brownish or tan. The ochraceous brown sides It seems callous to me, who is enchant- merge gradually to a whitish ventral fur. Pale ed by chipmunk charm, but I have known one facial stripes about the eye, above and below, man and another (as a youth) who each shot set off the prominent, bright black eye. The 200 chipmunks in and about their garden and dorsoventrally flattened tail, well-haired but not lawn for the sport of shooting, justified by bushy, is brown above with pale grayish bor- garden damage. I also knew of the Olsons in der and is dark chestnut or ochraceous bright Amherst Junction, who cut a hole in their below. There seems one molt per year (in the kitchen door to allow one of these delightful growing season) (Snyder, 1982). creatures to share meals. In 40 years, I have examined three live Although nowhere in danger of extirpa- melanistic chipmunks in Stevens Point, Wis- tion, the subspecies population of Tamias consin. All were released on my property, but striatus doorsiensis is vulnerable because it never seen again. James Woller caught a is found only on the Door Peninsula of Door melanistic male on 17 June 1995, in Stevens County. Point, which the author prepared (UW-SP Habitat. Eastern chipmunks are found in 8,016). Since there are thousands of eastern rocky places, such as bluffs or gullys, or on chipmunks in this community, the incidence exposed outcrops and ledges. They also thrive of melanism is certainly low. in hardwoods (oak, maple, hickory, and Measurements are given in Table Rod-3. beech), and in conifers especially in uplands Dental Formula. The dental formula is I where red and white pines grow in open for- 1/1, C 0/0, P 1/1, M 3/3 = 20. est. Along lakeshores, rivers, and at the edge Permanent dentition erupts by 3 months, of marshes and swamps they may be found, and yearlings have worn check teeth (Sny- and in jack pine savannahs and brushy hill- der, 1982). sides they are often abundant. They thrive in

TAXONOMIC ACCOUNTS / ORDER RODENTIA 181 human neighborhoods where trees and brush are present to provide food. Stumps, deadfalls, stone fences, and woodpiles provide perching places for them to defend feeding territories. They need soil depth because they must dig burrows to survive the winters. Being so well adapted to mature deciduous forests and to forest edge, Mahan and Yahner (1998) compared the demography of this species in continuous forests and mangaged clear-cut forest sectors. They found no differences in reproduction, body weight, time remaining on the study plots, or diet. When

Maps showing geographic distribution of Tamias striatus in Wisconsin and North America.

182 THE WILD MAMMALS OF WISCONSIN mast was scarce, both populations suffered York to prefer small acorns of white oak (i.e., the eastern chipmunks weighed less). In (Quercus alba) over red oak acorns (Q. ru- a related study (1996) burrow sites selected bra). Activity patterns of harvesting white oak on high ground, where logs and stumps acorns seemed to influence the success and seemed essential, were important as a “criti- development of the forest. They detect ripe cal” resource for these chipmunks. Surpris- and edible hazelnuts and cast off the husks to ingly, an overstory of trees, even oaks and eat them. It is essential a chipmunk hoards other mast producing species, was not so im- up seeds and acorns to last the winter, be- portant, because foods were varied. Such find- cause the winter sleep lasts only short peri- ings bode well for the eastern chipmunk; they ods of days and then the chipmunk emerges suggest that in clear-cut forest and suburbia, from its torpor and feeds. so long as den sites are available, chipmunks Snow lay deep in Stevens Point, several may thrive. Factors such as slope, stumps, inches at my bird feeder (25, 27 November perch sites, and diverse foods allow the chip- 1985), where an eastern chipmunk scrapped munks to select suitable den sites (Henderson with winter birds for seeds. A larger chipmunk et al., 1985). was seen foraging on snow nearby (24 No- Chipmunks do not always dig burrows, vember). I have seen them sunning on the but may live in nests in woodpiles and rocky snow-step of the burrow in early spring. In crevices. Burrows may be short or long and mild winters, such as 2000, they emerge in complex, probably depending on their age and central Wisconsin by 1 March; another was usefulness. Descriptions of burrows are given seen in Milwaukee the same date. by Panuska and Wade (1956). Much digging In Virginia, Wolff (1996) found high mast is carried out in winter, so that those burrows production to increase survival of Peromyscus are usually complex. In spring short burrows and chipmunks. This agrees with my obser- are built and often abandoned, as the chip- vations, but an abundance of chipmunks in munks fiercely fight and drive the losers out of fall is often followed by scarcity in spring. the area. Weaned young soon fight for space. Reproduction. The eastern chipmunk Foods. Eastern chipmunks feed on plants regularly produces two litters per summer. primarily, mostly acorns (mast) and maple Matings are frequent in late spring and seeds gleaned from the forest floor or pro- late summer. Testes regress in winter. The cured by climbing into the trees to harvest. young-of-the-year probably do not breed, but Very likely beech nuts are also eaten in beech they are observed trying at a tender age. Some forests in eastern Wisconsin. Other parts of tiny youngsters are seen playing about in late the plants are also eaten, buds, even the flow- summer. Judging from their size they likely ers (Aldous, 1941) but not so much as fruits, will never store up sufficient food for the win- nuts and seeds. In May, eastern chipmunks ter. After some winters, probably due to a ate mostly roots, according to Wrazen and shortage of acorns stored during the previ- Svendson (1978). Insects, bird eggs, inverte- ous autumn, hardly any eastern chipmunks brates, even mice and snakes may be eaten emerge from winter sleep. Those that do, (Torres, 1937; Harriot, 1940; Mumford and produce litters as early as late March. Males Whitaker, 1982). Fruits are eaten on the spot, are reproductive even in February, and occa- their seeds dropped on the ground, and seeds sionally come out to sun near their entrances and acorns are the most important foods in the snow cover. They probably occasional- cached underground for winter. Chipmunks, ly breed at this time. then, are very important in planting forests The females have two estrus cycles, in and ground cover (e.g., greenbriar). Pyare et the spring and summer (Banfield, 1974). The al. (1993) found eastern chipmunks in New gestation period is about 31 days, and the

TAXONOMIC ACCOUNTS / ORDER RODENTIA 183 young are born blind and naked, in the old approximately two acres. Often they do not winter burrow as a rule. By 18 days, hair is move about this much, depending on food evident on the skin. By a month their eyes resources, densities of the chipmunks, etc. have opened and they resemble small adults. Use-frequency is highest near the burrow, They then emerge from the burrow (after 5-6 which is central in most home ranges. Getty weeks) and play together. By five or six weeks (1981) found that chipmunks avoid one an- the mother recommences breeding. Her other allowing overlap of home ranges and young are gone when the second litter arrives spatial time-sharing. While winter hoards are a month later. Late-in-the-season litters, even collected, more time is spent fighting and emerging from their burrows in early Octo- underground. This “space-time” analysis de- ber, have only approximately a month to fines home range as time spent wandering, cache foods, but occasionally all the chip- separate from other activities. I can confirm munks have gone underground by then. (see Jackson, 1961) that chipmunks may at- The litter size is variable, usually 3 or 4 tain densities of 20 per acre, for they do that young, 4-5 according to Snyder (1982). The on my property regularly not even counting observed range is 1-8. There are eight mam- the newly emerged young. Burt (1940) re- mae. To my knowledge, the young never suck- ported about four per acre (10/ ha). le above ground, and possibly weaning forc- Remarks on Behavior. This small sciurid es them to emerge from the burrows. Tamias is diurnal, active from March (sometimes late lives up to 4-7 years (Crandall, 1964). February) to October, when it enters winter Mortality. The diurnal eastern chipmunk sleep. Torpor varies during the winter with fre- is seldom taken by owls. Hawks regularly take quent arousals, but Snyder (1982) says some them. I have seen a sharp-shinned hawk pick remain dormant all winter. There is no heavy one up and carry the poor little fellow away. deposition of fat in autumn (see Brenner, 1975; There are records of great blue herons and Scott and Fisher, 1972; Panuska and Wade, ravens feeding on them (Harlow et al., 1975; 1956; Maclean, 1981). Panuska (1959) shows Connor, 1971; Porter, 1951; Burt, 1941). the effect of warm temperature on “hiberna- Wild and domestic carnivores catch them, my tion” in Tamias, i.e., there is weight gain in own little beagle-terrier caught several in my cool burrows and significant weight loss in a yard. Raccoon, red fox, bobcat, lynx, coyote, warm environment. Rectal temperatures fall red squirrel, Norway rat, large snakes, and (from 38.50 C) with ambient temperature; as especially weasels and martens have killed torpidity is assumed, the rectal temperature them (Allen, 1938; Toner, 1956; Wood, approaches (to 30 C) the ambient temberature 1922; Banfield, 1974). (30 C) (Maclean, 1981). Snyder (1982) reviews Botflies (their larval warbles) infect chip- other physiological information. munks (Ozoga and Phillips, 1964; Domey, The animals are solitary and highly terri- 1965). They always seem to be scratching. torial except when a male and female are Fleas, lice, ticks and a variety of worms infest chasing about breeding. I have observed chas- these squirrels (Allen, 1938; Scharf and Stew- es even in trees at elevations well over 60 art, 1980; Timm, 1975; Snyder, 1982; and feet. Twice I saw chipmunk in McDill Pond, Jackson, 1961). The La Crosse encephalitus swimming across the channel to our island virus was identified in Wisconsin chipmunks (Maribell Isle), once making use of a sub- (Berry et al., 1975). merged log to get partly across the expanse Home Range and Density. Home range before entering the water. for eastern chipmunks varies from 100 m2 to Geographic Variation. The race T. s. gr- >10,000 m2 (Snyder, 1982). Males wander iseus differs from the peninsular race T. s. farther than females, and may range within doorsiensis in darker color, of the non-white

184 THE WILD MAMMALS OF WISCONSIN stripes and dorsal fur, of the feet, and of the the substrates reduced predation on those chip- underside of the tail. Jackson (1961) recog- munks having them. Some pale color in chip- nized a race in southern Wisconsin, referred munks of the Upper Peninsula ascribed to in- to T. s. ohionensis. He wrote that the chip- tergradation between T. s. peninsulae of Low- munks in southern Wisconsin, from Buffalo er Michigan and T. s. griseus may be ascribed County in the west to Washington County in to long ago introgression from populations the east were darker, more brownish, and leading to speciation of doorsiensis. Or it may smaller, but I noted no important distinctions. result from polymorphism in alleles for pallor, In Illinois, just to the south, Hoffmeister (1989) but strictly selected for, in their respective en- assigned northern specimens to the race T. vironments in both Michigan’s eastern penin- s. griseus, and did not recognize T. s. ohion- sulae and Wisconsin’s doorsiensis. ensis Bole and Moulthrop at all. There may be a slight clinal darkening in Jackson (1961) believed the pale chip- mainland Wisconsin chipmunks southward, munks on the Door belonged to the race T. s. but to follow Jackson (1961) assigning south- peninsulae Hooper, from Lower Michigan, but ern specimens to an Ohio race on such sub- Door chipmunks differ from that pale race in tle, gradual variation may add to confusion. wider cranial breadth, smaller skulls, and oth- Specimens from northern Illinois were as- er features mentioned by Long (1974). In cra- signed by Hoffmeister (1989) to griseus. I nial characters the Wisconsin chipmunks are recognize no significant speciation in south- all similar, suggesting the Door County chip- ern Wisconsin. munks became pale evolving locally on the pale limey soils and pale gray and buff limestone outcrops of the Niagara escarpment. Genes Tamias striatus doorsiensis Long for pale pelage may have been distributed widely, on the other hand, to be swamped out by 1961. Tamias striatus peninsulae Hooper, see Jack- the invasion from the south of darker T. s. son, Mammals of Wisconsin, p. 149. griseus. The alleles may have persisted on the 1971. Tamias striatus doorsiensis Long. Proc. Biol. Door Peninsula, where the substrate favors Soc. Washington, 84:201-202. Type from Pen- retention of the pale pelage. That would ac- insula State Park, Door County, Wisconsin. count for the close resemblance of the skulls of the Wisconsin races, and the marked size Description. This chipmunk is pale, not difference with small skulls from chipmunks to only its feet, cheeks, and dorsal pelage, but the eastward across Lake Michigan and be- also the underside of the tail. The rump patch yond. The short time since the glacial ice had is tawny and less rusty chestnut. The skull is covered the Door and filled Lake Michigan, similar to that of T. s. griseus. Compared to less than 10,000 years ago, does not allow T. s. peninsulae Hooper, from lower Michi- much time for either the evolution of T. s. gan and some of the Beaver Islands, a race doorsiensis responding to natural selection for which is comparatively as pale, doorsiensis has pale camouflage, or even the other possibility, much brighter, more conspicuous post-auric- of immigration. Long (1965) reported similar ular patches, more grayish dorsal pelage adja- post-Pleistocene differentiation of color and cent to the dark and white stripes, the tail is pallor of chipmunks and other sciurids in Wy- more frosted with white, more heavily inter- oming in the same span of time, which may mixed with black, and longer. The underside be comparable to what happened in Wiscon- of the tail is a shade paler than in peninsulae. sin. Probably the genes for pale pelage were The feet and cheeks are more ochraceous. The present in populations on both sides of Lake skull is more robust, with wider nasals, zygo- Michigan, and were rapidly selected for where mata, and rostrum. The teeth are more mas-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 185 sive, as in T. s. griseus. See account of the Specimens examined. Total, 73. Upper significantly darker Tamias striatus griseus. Michigan 9. Wisconsin. Adams, Burnett, Bay- Specimens examined. Total, 22. Door field, Brown, Chippewa, Clark, Columbia, Co.: Peninsula State Park 9 or more (USNM). Dane, Dodge, Douglas, Dunn, Forest, Grant, 6 mi. W Sturgeon Bay 4. Sturgeon Bay 3. 2 Iowa, Iron, Jefferson, Juneau, Kenosha, Ke- mi. N Jackson Port, near Wayside 2. Near waunee, Langlade, Lincoln, Manitowoc, Mar- Fish Creek 1. 3 mi. N Bailey Harbor on Hwy athon, Marinette, Menominee, Milwaukee, 57, 1. .5 mi. SE Peninsula Center 1. 1.5 mi. Oconto, Oneida, Ozaukee, Pierce, Portage, W Kangaroo Lake, on Hwy A, 1. Racine, Rock, Rusk, Sauk, Sawyer, Shawano, Sheboygan, Taylor, Trempealeau, Vernon, Vilas, Washburn, Washington, Waupaca, Tamias striatus griseus Mearns Waushara, Winnebago, Wood counties. Michigan Schoolcraft County. 1891. Tamias striatus griseus Mearns. Bull. Amer. Mus. Nat. Hist., 3:231. Type from Fort Snel- ling, Hennepin Co., Minnesota. Genus Tamiasciurus Trouessart 1942. Tamias striatus ohionensis Bole and Moulth- Douglas Squirrel, Chickaree or rop. Sci. Publ., Cleveland Mus. Nat. Hist., Red Squirrel 5(6):135. Type from Cincinatti, Ohio. Refer- ring to synonomy only Wisconsin specimens, “In the North, the red squirrels are... dependent on the but likely the race is invalid. seeds of various conifers... the refuse [from “feasts” at “feeding sites”] may form sizable midden heaps... The Description. Tamias striatus griseus is a red squirrel is partial to mushrooms and stores great small sciurid having striped pelage above and numbers [and] seems immune [even] to the toxic white below, thin whitish stripes on the eyelids qualities of the amanitas, eaten in some quantity, as this extending anteriorly to the nose and posteri- writer has had opportunity to observe”. — W. J. orly almost to the ear pinna. The median dor- Hamilton, Jr. American Mammals. sal stripe extends from the nape of the neck posteriorly onto the rump, but is not contin- Just a tawny glimmer, a dash of red and gray, ued to the base of the tail, the lateral stripes Was it a flitting shadow, or a sunbeam gone astray ! are likewise short, especially the lateral ones. I hear a mocking chuckle, then wrathful, he grows bold, Adjacent to the mid- dorsal stripe are narrow And stays his pressing business, to scold. — strips of brownish or grayish pelage, bordered Anna B. Comstock below with black, then comes white or creamy white, then black, then brown-ochraceous or grayish ochraceous. The feet are cinnamon brown, the tail grizzled grayish brown above and dark brownish orange below. Pale postauricular patches are behind the ear pinnae. The rump is rusty brown to dark chestnut. For comparison with the least chipmunk Eutamias, see account of that species. Com- pared to the pale race on the Door Peninsu- la, T. s. griseus has darker underside of tail, feet, cheeks, and the rump patch, and the upper side of the tail is less frosted with white. The cranium is similar. Red squirrel. Lloyd Sanford, with Hamilton, 1939.

186 THE WILD MAMMALS OF WISCONSIN Tamiasciurus hudsonicus (Erxleben) not bowed outward much, but are nearly par- Red Squirrel allel to the skull. There are eight mammae (one pair pectoral, two pairs abdominal, one 1777. [Sciurus vulgaris] hudsonicus Erxleben. Sys- pair inguinal). tema regni animalis..., 1:416. Hudson Bay. The color is typically reddish, with more Type locality fixed by A. H. Howell at Mouth or less golden tawny or grayish brown tones Severn River, Ontario, Proc. Biol. Soc. Wash- intermixed. The dorsal fur is lightly flecked ington, 49: 134, 1936. with black. In summer there is usually a bold, 1923. Tamiasciurus hudsonicus: Pocock. Proc. black line laterally and immediately above the Zool. Soc. London, p. 213. white underparts, extending from the shoulder to the flank. This lateral stripe is obscure The name Tamiasciurus uses the Lati- in winter. The head is darker than the dor- nized and Greek words meaning steward and sum, and around the eye is a conspicuous shade-tail and was doubtless intended by Po- white ring. The tail is bright reddish above, cock, with usage as late and as enlightened with the tips of the hair tawny and having as 1923, to point out the hybrid-like similar- subterminal black bands. The tail below is gray ities of this little tree squirrel to the other two and buff intermixed. The feet are tawny gray genera. Hudson Bay was the meaning of hud- or brownish gray with gray soles somewhat sonicus. The early naturalist Pennant de- overlain with silvery hairs. The underparts are scribed red squirrels from Hudson Bay. white or pale creamy buff. Description. Aside from its reddish color This reddish color, so much like the col- (see below) the red squirrel, which superficially oration of the red fox Vulpes, may cause in- resembles the Sciurus, has distinctive fore- timidation of the other tree squirrels, which feet and genitalia and is smaller in size. The are often set upon by even the young red squir- forefeet, adapted both to spending much time rels. In most of the western subspecies of this on the ground and to leaping about in the squirrel, the dorsal fur is dark tawny brown tree canopies (mostly horizontal branches) or even gray-brown, the tail tipped with con- have inner toes (III, IV) nearly equal in length, spicuous silver-tipped hairs as in the gray squir- and those adjacent (II, V) also are subequal rel. For this reason, westerners are not likely (Long and Captain, 1974). The thumb (I) pos- to call the squirrel a red squirrel, and often sesses a minute nail as in Sciurus. Some have use the name chickaree (Long, 1965; Hall referred to the even arrangement in Tamias- and Kelson, 1959: 399). ciurus and Eutamias as artiodactyl, which is The coloration of red squirrels in Wis- somewhat misleading because there are, of consin boggles the mind at times, because the course, an odd number of toes (five). variation in one place may include some par- The genitalia are unusual in both sexes ticularly brown, others having reddish mid- compared to those in Sciurus. In males there dorsal bands, and others being quite rufes- are minute Cowper’s glands, no bulbar gland, cent. When the mid-dorsal band is present, a true urethral diverticulum, a long filiform the sides are buffy or cinnamon brown. This penis, and a minute os penis. The seminifer- marked variation is seasonal, and usually re- ous vesicles are huge. In the female the vagi- lates to the molt. The molt apparently begins, na is unusually long and coiled during estrus. especially in young-of-the-year, in the tail with Anal scent glands are present (Mossman et a sketchy extension of red developing along al., 1932; Layne 1954, in New York squir- the back. This molt takes place in fall, and by rels). The tail is about half the length of the November the mid-dorsal band is common in body, the ears are somewhat pointed and tuft- the squirrels with sides and feet pale golden ed (especially in winter). The zygomata are brown or cinnamon gray. The winter pelage

TAXONOMIC ACCOUNTS / ORDER RODENTIA 187 is much brighter red than in summer, although Status. The red squirrel is holding its own the feet are often grayish in winter. The molt within its distributional area, especially in co- seems to progress forward from the tail (No- niferous forests, but not so well in urbanized vember) and subsequently spreads laterally and areas or other areas of hardwoods densely onto the shoulders and head. The bright hairs populated by gray squirrels (Reichard, 1976; seem to fade, not molt, after April, when the Riege, 1990). It avoids treeless grasslands and fur is worn and more brownish. marshes, but is a conspicuous denizen of the See Table Rod-4 for measurements for northern fir, cedar and pine forests. It has lost the two sexes. The size is significantly smaller ground in the extreme southeastern part of than in Sciurus. Wisconsin and in adjacent counties of Illinois Dental formula. I 1/1, C 0/0, P 1-2/1, (Hoffmeister 1989). m 3/3 = 20-22. The anterior (third) premo- Red squirrels in the United States are lar of each upper jaw is usually absent, and if seldom hunted or harvested for fur. Indians present minute, and partially covered by the used them for food (Cleland 1966), and some large fourth premolar. people in central Wisconsin eat them togeth- Geographic Distribution. Widely distrib- er with gray squirrels. In Canada 25,000 skins uted in Wisconsin, except in the southwest- are sold annually (Banfield 1974) to fur trad- ern part of the state. Surprisingly it is absent ers and subsequently shipped to Europe. They southward of the Wisconsin River from Co- have a value of only two or three dollars each. lumbia County, westward to the Mississippi. Red squirrels are not protected by game laws. There are no records in northwest Illi- They are responsible for some tree damage nois, although the area is ascribed to the prob- by gnawing, eat thousands of pine seeds, kill able range of this species by Hoffmeister young birds, and eat the eggs of songbirds on (1989). This squirrel is another example of a occasion. Some mistaken squirrel hunters Jackson-Hoffmeister pattern (see Plan of this have the belief that their fighting with (and Book, Plan-1), but the zoogeography is in- the supposed castration of) the larger Sciu- deed more interesting than that simplistic tax- rus is bad for squirrel hunting. onomy implies. Apparently the Minnesota Red squirrels have endearing aesthetic race has worked its way into Wisconsin from qualities, being such a beautiful and conspic- the west and northwest, consistent with a uous member of the North Woods. Fishing pattern of other forest mammals of western with my two sons some years back, on a Ca- origins. There it intergrades with the eastern nadian lake, we recall red squirrels as memo- race T. h. loquax (Bangs). rable as the loon by the boat, the bald eagle that soared overhead, the osprey, and the lines of mergansers fleeing and pattering over the water. It seems now more remarkable than any of the fish we caught. Where the red pines grew tall on high granite outcroppings tower- ing over the shoreline, red squirrels were leaping about some 150 feet (50 m) above our boat. The squirrels seemed so small at that distance, and leaped so far from tree to tree that they seemed like little birds. They never once fell. I have not seen any such “frolick- ing” in tree canopies in Wisconsin. Habitat. This squirrel is found common- Skull of Tamiasciurus hudsonicus. ly in northern forests (Hatt 1929, Layne

188 THE WILD MAMMALS OF WISCONSIN 1954), usually in pines, balsam fir, spruce, hemlock, cedar, and mixed hardwood-conifer forest (with beech and maples). The species may be found on occasion where the conifers are introduced and the typical trees are hardwoods (oak, hickory). In southeast Minnesota red squirrels are found in several towns. On the campus of St. Olaf College, in Northfield, Minnesota, the habitat is hardly coniferous, but a few red squirrels range around the edge of the campus. The red squirrels in Iowa live in pure deciduous tree stands near rivers and lakes (Lynch and Folk 1968). Sometimes in Wisconsin the red squirrel is

Maps showing geographic distribution of the red squirrel in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 189 found in jack-pine-red oak savanna or in red cy 1980, Layne 1954, Doutt et al., 1966). A pine and spruce Christmas tree plantations. tree nest is usually the approximate size of a They are abundant in parks where white pines basketball, with twigs outside a layer of leaves. are predominant. In beech-maple, with some Inside is the nest material of fine vegetation, hemlock or pine intermixed, either red squir- usually grasses. I have noticed a mother and rels or grays are predominant. This is true her young carrying grasses into a hollow oak also in lower Michigan, where fox squirrels tree formerly occupied by white-breasted prefer the oak and hickory (Beckwith 1954). nuthatches. In the cedar siding of my porch, One can say the same for southwestern Wis- there was no nest material at all for the young consin, where ecological preference may at squirrels, suggesting that this den was tem- least back up the factor of the river barrier porary. It was used at least twice, and at least maintaining reds north of the Wisconsin Riv- for two days (Long, 1993). er and excluding them southward. In these The mother accompanies her young reg- river bottoms, sycamore, hickory, even wal- ularly from den to den, and some dens con- nut trees forest the area. Hemlock, pine, tain more nest materials than others. When spruce and fir are not common, and there is young are discovered or threatened the moth- little of the preferred habitat for Tamiasciu- er presents an intense distraction display and rus. I found huge stands of jack pines. Since afterward carries the young, one at a time, by red squirrels are active in winter, it seems they its ventral skin, away to a safer den (Long, would readily cross the ice wherever the Wis- 1993). Temporary dens for young have been consin River is narrow and the current not called “outliers” following the lore of the Eu- swift. Possibly red squirrels only recently ap- ropean badger (Long and Killingley 1983), proached the river, and are not really com- these differing in their permanence, occupan- mon in river bottoms, which may, along with cy by other members of the social group, etc. the river itself, act as an effective barrier. There is similarity between movements of On islands in Lake Superior and Lake young to several outliers in these two quite Michigan, red squirrels are more often found different species: the arboreal or semi-arbo- than any other squirrel, usually exclusive of real red squirrel, generally being rather soli- any other kind of squirrel. On these islands tary, and the fossorial European badger, fa- maples and cedars are abundant, and there mous for its complex burrow systems and are some oaks, pines and fir trees. On Stock- social clans. ton Island, in Lake Superior, William McKee In eastern states, red squirrel territories (field notes) collected them in mixed forest defended by adult male or female were not (white and yellow birch, maple, cedar, hem- common as observed in this species in west- lock, and balsam fir). On Washington Island, ern states (Riege, 1990; Layne, 1989; and in Lake Michigan, red squirrels are seldom seen others). Red squirrels often defend against far from balsam and hemlock. In the southern conspecifics (Hazard, 1960). Where the gray parts of the range, red squirrels occur in quak- squirrels are abundant (Riege, 1991), even ing aspen, jack pine, white birch, hazelnut, and though the red squirrels are aggressive, they red maple. This habitat was north of New Lis- are usually not predominant. In late summer bon (field notes, L. Wargowski). there may be competition for tree cavities The denning of Tamiasciurus is fitting (Long, 1993). for its hybridized name, for they den in tree Foods. In central Wisconsin, red squir- cavities and leaf nests in trees as the Sciurus rels were observed feeding on jack-pine cones do, and they den at ground level even in bur- of both sexes, flicker eggs, fruit-heads of sweet rows (Yahner 1980), and also in woodpiles fern Comptonia sp, and hazel nuts. The red and in jumbles of rock, as chipmunks do (Fan- squirrel may feed on any seeds or fruits in

190 THE WILD MAMMALS OF WISCONSIN season. These include acorns, conifer seeds, Internal parasites include tapeworms, flukes, nuts (Hatt 192, fungi and cambium bark roundworms, bot fly warbles, and coccidea (Smith 1968, 1981), buds, flowers and ma- (Dorney 1965, Baker 1983). ple seeds (Reichard 1976) and sap of trees, Reproduction. Breeding begins February raspberries, insects, young birds and small and March and re-commences in June and mammals (Klugh 1927, Hamilton 1939, Hat- July. A female from Jackson County was lac- field 1937, Kilhain 1958, Banfield 1974). tating as late as October 10. Several males Smith (1981) measured food consumption as may attempt to mate one female. Layne (I954) about 117 kcal per day, for a lactating fe- believed that there were attempts at monog- male 323 kcal per day. See Table Rod-1. amous mating, when a male included a fe- In late summer and fall the red squirrels male in its home range and defended the area begin to cache food for winter, mostly seeds, to keep other males out. There are soft cough- nuts, and acorns. In some places the food ing calls and chasing of the female prior to stores are cached in “middens” where hun- copulation in a tree or on the ground. Re- dreds of food items are stored. Most charac- ported gestation periods range from 34 to teristic in middens I have observed are stores 45 days (Ferron and Prescott, 1977; Lair, of conifer cones. These may be buried, or 1985). Litter size is about 5 (4-7). The young laid on the ground at the foot of a tree. Re- are born hairless with eyes and ears closed. portedly, the tree as “prey” and the squirrel They are about 70 mm in length, and weigh as “seed predator” have co-evolved with mu- about 6.5 grams each. Hair develops by 10 tual benefit (Smith 1970, Elliot 1974). In days, and the eyes open in about four to five spring, when chasing or calling, the red squir- weeks. The young venture from the nest by rels occasionally fed on cached cones. In sum- 38 days, and are weaned by 7 or 8 weeks mer, in central Wisconsin, I saw them simul- (Ferron 1980, Layne 1954, Svihla 1930). The taneously calling from tree canopies and feed- young accompany the mother learning, playing on jack-pine cones. During May-June red ing, finding things to eat; they may stay with squirrels foraged chiefly on red maple seeds, the mother all winter on occasion. This ma- according to Riege (1991). Where and when ternal behavior suggests that usually only a available, red squirrels feed on maple seeds single litter is brought forth in a season, but in Wisconsin. there are usually two litters. Young do not Mortality. Humans are an important en- breed in their first year. One squirrel lived emy, shooting red squirrels and killing them three years in the wild, and they may live 10 by automobiles. Forest fires cause mortality. in captivity (Banfield 1974). The marten is a famous predator (Seton Home Range and Density. Riege (1991) 1953). The incredible climbing abilities of the found evidence of a “spring shuffle” when red red squirrels certainly lessen predation by ar- squirrels move about freely and their home rang- boreal pursuers. Predators on red squirrels es overlap. In winter red squirrels avoid the hard- include badger, bobcat, red fox, screech, woods and their home ranges diminish. barred, and barn owls, great horned owl, red- Density is difficult to measure as the squir- shouldered hawk, Cooper’s hawk, broad- rels move about so much. Layne (1954) found winged hawk, coyote, mink, long-tailed wea- the density to vary with the habitat quality. sel, lynx, kestrel, and snakes (see Baker 1983). For his study, one for five acres was low (1 / Parasites include fleas, 11 species of fleas 2 ha), and two per acre (10 / 2 ha) high. in New York alone (Layne 1954), and two in Food concentrations and family gatherings Michigan (Monopsyllus and Orchopeas, may raise density to 12 per acre, according Scharf and Stewart 1980). Lice, ticks, mites to Jackson (1961). I have seen as many as and chiggers are reported from red squirrels. seven per acre in jack pine savanna, but five

TAXONOMIC ACCOUNTS / ORDER RODENTIA 191 were offspring of one female. The males usu- ward into Illinois and eastward into Indiana. ally come and go, ranging out of the one- Without specimens from southeast Wisconsin acre study area. I have seen red squirrels ac- to examine, except one from Germantown, tively chasing one another in late August on this assignment is purely on geographic Washington Island, doubtless territorially grounds, a line drawn subjectively between to claiming some foraging area for winter food distinctive samples. From Table Rod-4, I judge supply. Rusch and Reeder (1978) felt that ter- Hoffmeister (1989) was correct in saying T. h. ritorial behavior regulates red squirrel density loquax has larger feet. Jackson’s comment near pine cone sources. Smith (1968) sug- (1961) that minnesota had larger feet, by 10 gests that red squirrel males expand their feed- %, is in error, although some individuals may ing territories in spring to include females, be large. The geography is set forth below. while chasing off other males. Riege (1991) found greatest densities are in autumn. Lin- duska (1950) found 50 percent turnover in Tamiasciurus hudsonicus loquax (Bangs) red squirrels. Remarks. Long (1993) recorded several 1896. Sciurus hudsonicus loquax Bangs. Proc. Biol. interesting behaviors of the red squirrel. Used Soc. Washington 10:161, type from Liberty as a nest-site distraction display, and on oc- Hill, Conneticut. casion (but with less intensity) in territorial and 1936. Tamiasciurus hudsonicus loquax: A. H. interspecific threats, a complex call involving Howell. Occas. Papers Mus Zool., Univ. Mich- two or more vocalizations practically at once igan. 338:1. Specimens examined. None. is emitted in a conspicuous display by a mother Hoffmeister’s assignment, based on Cory red squirrel near young. This call involves, with (1912:124). the harsh chattering, a “piping” or squeaking component resembling young. Along with the mother’s leading of the observer or predator Tamiasciurus hudsonicus minnesota from tree to tree, it distracts attention away (J. A. Allen) from the young hidden in a nearby nest. The same call was often given at dawn by the moth- 1899. Sciurus hudsonicus J. A. Allen Amer. Nat., er, sited a short distance away from an outlier 33:640, type from Fort Snelling; Hennepin nest from which the young quietly emerged. Co., Minnesota. There is a ventriloquistic quality to the call, 1940. Tamiasciurus hudsonicus minnesota: Hay- which probably confuses the observer about man and Holt. In Ellermann, The families and the call’s direction and source. The summer genera of living rodents. British Mus., 1:346. movements of the mother and her young from 1943. Tamiasciurus hudsonicus murii A. H. How- one outlier to another in the family’s wander- ell. Proc. Biol. Soc. Washington 56: 67, type ing and feeding, sometimes changing dens from Morehead, Clay Co., Minnesota. from night to night, are quite interesting. Additional natural history. Steele (1998) Specimens examined. Total, 143. Ad- reviewed the biology of the red squirrel. ams, Ashland, Bayfield, Brown, Calument, Geographic variation. There are two sub- Chippewa, Clark, Door, Florence, Forest, species of red squirrels ascribed to Wiscon- Iron, Jackson, Kewaunee, Langlade, Lincoln, sin. Throughout most of the state and in Up- Manitowoc, Marathon, Marinette, Oconto, per Michigan as well, the race is T. h. minne- Oneida, Outagamie, Portage, Price, Sauk, sota. Interbreeding in southeastern Wiscon- Sawyer, Sheboygan, Taylor, Vilas, Wash- sin, the southeasternrnmost specimens are burn, Washington, Waukesha, Waupaca, ascribed to T. h. loquax, which ranges south- Waushara, Wood counties.

192 THE WILD MAMMALS OF WISCONSIN Table Rod-4Rod-4. Measurements of 14 males and 6 females Sciurus carolinensis Gmelin of Tamiasciurus from Portage County, Wisconsin. Gray Squirrel TL Tail Hind foot Ear Males 308.8±10.8 125.6±5 44.9±1 19.4±3 1788. [Sciurus] carolinensis Gmelin. Systema naturae, Females 317±21 125.2±8.5 43.0±5 19.3±4 ed. 13. 1: 148. Type locality, “Carolina.” Greatest Zygomatic Interorb. Max. Length breadth breadth t-r Sciurus carolinensis pennsylvanicus Ord Males 45.8±1 26.2±0.8 13.9±.5 7.5 ±.26 Females 45.4±0.6 26.1±1.1 14.1±0.6 7.8±0.3 1815. Sciurus Pennsylvanicus Ord. In Guthrie, A new geog., hist., coml. grammar... Philadel- phia, 2: 292. Type from Pennsylvania, west of the Allegheny Mountains. Genus Sciurus Linnaeus 1894. Sciurus carolinensis pennsylvanicus: Rhoads. Tree Squirrels Appendix of a reprint by Ord, on page 19. 1961. Sciurus carolinensis hypophaeus Merriam: Medium-sized (see Table Rod-5) diurnal squir- Also Jackson (1961). Wisconsin. rels with long, bushy tails, prominent ears (usu- 1948. Sciurus carolinensis hypophaeus Merriam: ally tufted in winter, in Wisconsin), tiny pollex Burt (1948). Upper Michigan. bearing a minute nail or hoof distally, upper 1912. Sciurus leucotis: Cory. Field Museum Nat. third premolar vestigial or absent, digits IV and Hist., Zoology, Vol. XI, p. 116. Also Snyder, V elongate, upper molars showing prominent Bull. Wis. Nat. Hist. Soc., 1902. transverse crests confluent on lingual border, and often terminating in distinct cusps. Brain- The name Sciurus means shade-tail in case depressed posteriorly (i.e., cranium Greek, but it came to mean squirrel, which is arched), palate broad, zygomata broaden pos- how Linnaeus used the term. The specific terior to the upper toothrows, occiput extends name of the binomen, of course, refers to posterior to exoccipital condyles, often with a the type locality Carolina, and pennsylvani- median and minute projection in adults. cus refers to the type locality of this wide- ranging geographic race. Description. Generally the description is the same as given for the genus above, but in comparison with the fox and red squirrels, more trenchant characters are listed. The gray squirrel resembles the slightly larger fox squirrel, but is larger than the red squirrels (see measurements). The skull differs in smaller size from that of the fox squirrel. Al- most always there is a small peg upper third incisor present. In some states about 1% lack the peg incisor, but Jackson (1961) observed no specimens and I have only a few (for examples, UW-SP 134, 4963). The absences were observed only in subadults (having the fourth upper premolar not fully erupted). The baculum is small, disk-like with a thorny pro- Sketch of a gray squirrel Sciurus carolinensis. Courtesy jection (Burt, 1960). Chromosomes number Dan Metz. Scratchboard. 2N = 40 (Nadler and Sutton, 1967). There

TAXONOMIC ACCOUNTS / ORDER RODENTIA 193 are eight mammae (four pectoral, two ab- yellow and the head nearly white (UW-SP dominal and two inguinal). 6970, 1111). Over a third of a century I have The red squirrels and gray squirrels are observed one to a few yellow mutant gray the only Wisconsin tree squirrels with long squirrels in nearly every year. These mutants whitish hairs in the tail, but there is no trace occur in an area about 7 by 3 miles, on the of rufous in the fur of S. carolinensis, so char- east side of the River. Some squirrels show- acteristic of the dorsal fur of the red. The belly ing intermediate dorsal color seem heterozy- of the gray squirrel may be whitish, as in the gous for yellow and normal, and some of these red squirrel, but it is never whitish in the fox supposed “hybrids” are nearly gray with yel- squirrel. The light-colored venter in the gray lowish tails. In 1997, Professor Mark Boyce squirrel may vary, to be rusty, cinnamon, or told me he saw a yellow mutant on the west even brown. Often the belly is marked with side of the River, the first such report. In De- creamy or rusty blotches or a mid-ventral line cember 1999, my wife and I observed three of brown-tinged gray. The gray squirrel has yellows together chasing one another at the grizzled dorsal fur, but the predominant tone cemetery on Patch Street, near McDill Pond. of it is gray. Often in winter the dorsum is In late January and February, staying until mid- more or less frosted white. On the flanks there dle May, a mixed yellow (the tail certainly yel- is generally a brownish tone, and in winter low) was resident on our acreage, with one and early spring the ears are somewhat tuft- black and eight normals. ed with orange and white patches. This fea- In the melanistic squirrels there are ear ture seems for the most part to have been tufts present in winter, but they are obscure, overlooked; I have never seen tufts on the concolor with the dorsal fur. If there is an ears, even in winter, on gray squirrels from iron-gray brownish tone on the face, throat central Illinois or at Northfield, Minnesota. and chest of a gray, it also tinges the tufts. Tufts are variable in frequency in grays from The ear tufts are conspicuously white on the southern Wisconsin. yellow squirrels. Melanism (e.g., UW-SP 4042, 988-9, The molt commences in late spring, and 6968, 895, 3585) is common in the Wiscon- ear tufts disappear then (by late May). Molt sin populations (see below), and I have seen a continues into summer (Brown and Yeager, live albino on Washington Island. In Stevens 1945). The molt proceeds from the top of Point, which is incorporated on both sides of the head and mid-dorsum, and proceeds in the Wisconsin River, there is a creamy or yel- all directions especially posteriorly and ven- lowish mutant squirrel, the tail nearly lemon trally until complete. A second molt appears on the flanks in July-September and proceeds dorsally and anteriorly. This primary molt probably and almost cryptically begins in July (Flyger and Gates, 1982). Students often are confused by identification of gray squirrels because many specimens are not the ashy or silvery gray so common in late autumn and winter. This time is peak abundance of squirrels, the time they are hunted, and the time when leaves are down making it easier to see the squirrels. If gray at all in summer, the dorsum is somewhat rusty iron gray above, and often the belly is rusty, almost chestnut Skull of gray squirrel Sciurus carolinensis. brown. In most specimens the belly is pure

194 THE WILD MAMMALS OF WISCONSIN white, although the throat and upper chest may be yellowish tawny or rusty brown. From Washington Island one specimen had a rusty brown belly, and some venters observed from various localities in Wisconsin were more or less brownish or ochraceous- brown in other parts of Wisconsin too. The feet are often rusty and bright, and rusty brown hairs are found intermixed in the tail. New fur is short and brownish, about the same color as a snowshoe hare, and such a color does not fit expectations when persons are more familiar with the ashy gray. The long,

Maps showing geographic distribution of Sciurus carolinensis in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 195 whitish hairs intermixed with gray and brown rel. The great appreciation for this cute ac- in the tail is important in identifying brownish tive, intelligent and pretty mammal scamper- gray squirrels, especially in summer. These ing about in our lawns, parks, and natural are concentrated laterally in the tail, almost areas cannot be measured. It is often hunted as whitish outer bands. The ashy color of the for sport for its delicious flesh, not so much dorsal fur is a whitish tipping of the shorter for its fur. brownish fur becoming intermixed with long The gray squirrel occasionally kills young black ones as they grow out. The dark hairs song birds or eats their eggs, raids an occa- appear on the flanks and shoulders, and be- sional garden or gnaws down saplings and come confluent laterally and sometimes above small fruit trees, and gnaws on houses (one a persisting band or stripe of brownish gray. caused several hundred dollars damage to my Eventually new ashy fur spreads upward and deck drawn to the wood probably by the pres- toward head and tail. In summer, the short ence of sap in the cedar). Grays are so clever fur on the face is quite brown, and this color at robbing bird feeders some frustrated bird persists in some squirrels all year. The ear lovers are unhappy. [My wife hangs an emp- tufts grow out in late fall and winter. Measure- ty plastic milk bottle (appropriately trimmed) ments and weights are given in Table Rod-5. around the post, and it seems to always keep The grays are smaller than fox squirrels aver- the squirrels away, so long as they cannot aging 18-21 inches in length (457-553 mm) jump onto the feeder from above. Therefore, in total length, and up to 28 ounces. they forage at the base of the feeder becom- Dental formula: I 1/1, C 0/0, P 2/1, M ing fat and almost square in form.] One of 3/3 = 22. the most curious mysteries, now attributed to Geographic range. Populations of gray the gray squirrel, was the disappearance of squirrels occasionally increase into the thou- the tiny American flags put out in an Osh Kosh sands, forcing them in low mast years into cemetery over the Memorial Day and July 4th starvation or mass emigrations. These phe- holidays. For two years a thief was suspected nomena of high density and emigration were of stealing flags. When a strong wind knocked observed often in early Wisconsin history down a big tree, shredded flags were found (Schorger, 1949), prior to game magagement. within gray squirrel nests. The positives greatly The numbers of gray squirrels rose and fell outweigh the negatives, regarding the status cyclically, perhaps approximately every five of the gray squirrel, and being such a beauti- years (Jackson, 1961), and they varied then ful object of nature study and photography and do now considerably from place to place we are happy to have it here. and year to year. The number of acorns pro- Habitats. In a word, the habitat of the duced probably play a role in these local fluc- gray squirrel is “hardwoods”. The oaks and tuations (Nixon et al., 1975). Gray squirrels, maples are essential not only for seeds and to cache food, hide acorns in holes dug in the security from predators, but for the tree hol- ground, usually one per hole, never more than lows that often shelter the squirrels in winter a few. Therefore, the squirrels plant their own and provide security in the care of young. The forests and create their own habitat. Grays habitat is often called the forest edge, or open return to their caches, apparently by smell- hardwoods, overlapping the similar habitat of ing, reportedly finding as many as 85 per- Marmota and Sciurus niger. In Illinois for- cent of the hidden acorns (Thompson and ests, elective logging depressed breeding in Thompson, 1980). The surplus obviously re- old adults temporarily, but squirrels returned mains as seed; the gray squirrels are benefi- more abundant than ever (Nixon et al., 1980). cial to the forestry in Wisconsin. There is im- Riege (1991) found that availability of foods mense aesthetic value of this beautiful squir- and habitat affected densities of gray squir-

196 THE WILD MAMMALS OF WISCONSIN rels more than interspecific competition with roadway, the numbers vary from six to at red squirrels in the forests. Gray squirrels at- least 22. There are seven or eight old red tained their highest densities in oak-maple oaks, a few small white oaks and two large forests, whereas red squirrels concentrated, ones, a grove of hazelnuts, and there are for the most part, in fir-cedar communities or numerous maples present. There are also pines, when they were producing cones. Clear four other kinds of squirrels commonly cutting and clearing of trees improve habitats present, and three of these also feed on for grays (and other squirrels). When nest cav- acorns and maple seeds. When squirrels are ities are in short supply two or several gray numerous, there are movements of at least squirrels occupy the same tree cavity. Leaf one, sometimes several squirrels, to and from nests are also constructed, in oaks and other other neighborhoods. In a normal year, af- high trees, occasionally in pines. ter a mild winter, I counted as many as sev- One may evaluate habitat suitability in en squirrels chasing one another in a tree in the gray squirrel by reference to trees espe- April, with two others on the ground. Such cially those with acorns and other hard mast. chasing in similar numbers was also seen in In Wisconsin there are two chief cover types, mid-February. Some squirrels, known by tail namely deciduous forests (wetlands, high- damage or mutant color, wander at least a lands and urban parklands) and mixed de- kilometer away. ciduous-conifer forest (mostly in northern There is a British word for squirrel nest, counties). Especially in winter there must be the “drey”, used for their “grey squirrel”. hard mast, hoarded by gray squirrels and also Wisconsin gray squirrels often build nests of used by their competitors: chipmunks, white- sticks and leaves high in the tree canopies. footed mice, and others. They also carry vegetation into tree cavities. I For survival of the gray squirrels, there suppose the word drey applies to both kinds must be adequate cover and trees for secu- of squirrel nests. The nest for the first litter rity, for nesting in season, and for repro- and for most adult grays, when hollows are duction. Forests of large diameter trees hav- available to them, is usually placed in a tree ing dense overstory and numerous tree cav- cavity or some hollow structure similar to it. ities, and including some oaks provide opti- The nest often is lined with shredded bark mal habitat. Canopy closure varies from and plant fibers. Later in the year, the nests about half to 75 percent, and understory constructed in cavities may be bundles of should vary from 20-30 %. shredded leaves. Nest construction likely is There are two major requirements nec- somewhat opportunistic in site selection and essary for squirrels, and they may considered nest materials. Young squirrels and those not one. Oak trees, maples, or other food trees finding hollows are often forced to build leaf are necessary, and if winter is severe the nests, in the tree branches. These are made available mast food cached mostly in the clumsily by the beginners. In the best ball-like ground must keep the squirrels alive until squirrel nests, the intertwining of branches and spring. Therefore, bird feeders and other al- leaves makes a rain-proof chamber (entered ternate food sources benefit squirrels in ur- by the squirrel from an obscure opening at ban areas, especially those having some ma- one side) (Jackson, 1961). Sometimes these ture hardwoods. Hoffmeister (1989) found leaf nests are used for a couple years, and two mature oaks on the University of Illinois tree hollows perhaps as long as opportunity campus to provide for about five gray squir- exists. Leaf nests are usually sited about 25 rels, and the numbers varied from year to feet above the ground in hardwood trees, year. On my property, in central Wisconsin, but occasionally are built lower. Rarely gray about an acre bordered by lakeshore and squirrel nests are found in conifers. Occa-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 197 sionally the nest is built on an older squirrel breeding commences in January and Febru- nest or on a bird’s nest. ary, with young appearing after a gestation Foods. Jackson’s (1961) list of foods is of some 44 days. In 1968, when the snow in close agreement with foods that I have was piled high around my house some naked seen or heard that gray squirrels use, as fol- young fell out of a gray squirrel’s nest (sited lows: many nuts including acorns, seeds, in a jack-pine) in darkness right onto the ice, fruits, especially seedy fruits of maple, elm, and with a great commotion the mother car- hornbeam, hackberry, arrowwood and oth- ried them away somewhere in the darkness. er species of Viburnum, cherry, mulberry, First litters are usually born by early April, and thornapple, wild grapes, buds of elm and oak, breeding continues into July or later for sec- inner bark and sap (maple, elm), corn (the ond litters. In September 1994, I saw tiny germ only), insects, and occasionally bird young that emerged from the nest, in a bur eggs (Dambach, 1942; Korschgen, 1981; oak on the University campus. As many as Nichols, 1958; Montgomery et al.1975; eight young may be born, but usually the num- Sanderson et al, 1980). The gray squirrel ber is two to four. Only the mother cares for occasionally preys on small birds and mam- the young, and they may be moved if the nest mals, even an eastern chipmunk. Squirrels is disturbed (Nixon et al., 1968). Weighing gnaw on bones for the minerals. I have seen about 15 grams at birth, the young slowly them eating mushrooms and suet. In winter grow and fur out, opening their eyes at ap- and spring, gray squirrels make numerous proximately 30-35 days. In this museum col- excavations in the snow and ground in search lection there are several adults marked as lac- of buried acorns, usually carrying a wet acorn tating when collected, in early April, and one into a nearby tree to eat. I found them to (UW-SP 912) as early as 29 March. Weaning feed on oak buds in spring, hazelnuts in sum- begins after about 50 days when the active mer, and apples in fall. young are about half grown. By then the Some buried acorns are eaten after the mother is ready for her second litter. These snow melts, and I suspect that many acorns may stay with the mother into winter. One are hoarded and eaten in tree hollows. When small male (UW-SP 5) taken April 29, is juve- grays are fighting in winter they may be com- nile. Its greatest length of skull was only 45.3 peting for females, but also defending tree mm. The only teeth erupted were the inci- hollows used as homes and food caches. Ter- sors and the deciduous fourth premolars. In a ritoriality may be defense of area, the female season a mother usually produces at least six mate (as in deer), or a cache of acorns. C. young in the two litters, and in Longley’s study Smith (1978) suggests “spreading out food (1963) almost half the young survived through by scatter-hoarding avoids an effective, ag- one winter until the next autumn. Females gressive defense of a stored food site from are reproductive for about six years, and males one’s conspecific competitors.” Gray squir- for about eight. rels may scatter hoard nuts to reduce site ter- Mortality. By hunting and removing hab- ritoriality, whereas red squirrels that both scat- itat, humans are enemies of gray squirrels, ter hoard and hoard seeds in larders may not and automobiles kill them in Wisconsin successfully defend their caches from gray doubtless in many thousands. Add to human- squirrels, making the commoner gray squir- caused mortality many squirrels killed by rel a kind of parasite (parasitoid?) on the red dogs, cats, and electric wires. Nevertheless, squirrel. the species is holding its own today, espe- Reproduction. Some mother gray squir- cially in suburban habitats. By conserving old rels have two litters per year. They usually mast trees in lawns and parks, where hunt- cannot breed until their second winter. The ing is seldom, humans are not adverse to

198 THE WILD MAMMALS OF WISCONSIN gray squirrels. In nature bobcats (if present), less than 20 ha. Males had somewhat larger weasels, martens (when present), wolves (if home range than females. The home range present), red and gray foxes, and birds of varied inversely with squirrel density. Territo- prey kill them. They are not easily taken as riality was not evident. Madson (1978) report- prey. Ravens, snakes, and owls occasionally ed that 21 grays in Sauk County, Wisconsin, kill them. Once in Stevens Point, in winter, I wandered approximately 2 acres (0.8 ha). watched a gray squirrel on the side of a tree Dispersal was noted from August to Jan- directly facing a goshawk, which made an uary, but in some years it began as early as occasional pass, usually sitting on a nearby March. Where oaks were producing the few- branch within two feet of the bold squirrel, est acorns gray squirrels there had their low- but it could not catch it. The squirrel always est densities. Longley (1963) found home escaped by darting behind the tree. In early range (foraging) to extend as far as 5 miles (8 November (2003) a small Cooper’s hawk km). Live-trapped squirrels returned home alighted on a tree branch above a gray, cut- from distances of about 3 miles (= 4.8 km) ting off its escape, to force it to leap down (Schwartz and Schwartz, 1981). to the open ground. The gray immediately In large forest tracts grays seldom exceed ascended and bit at the hawk, driving it away. two per acre. Densities varied from one per In 25 years at our home, we have seen three acre in September to only 0.65 (1 per 5 ha) gray squirrels with bobbed tails (from fight- in winter. Baker (1983) reported 5/ha as gen- ing?). Some works on predation include eral. The high densities found years ago, when Schofield (1960), Pettingill (1976), Harlow 20 could be shot in a single tree, cannot exist et al. (1975), and Baker (1983). in the diversity of habitats today, according Parasites include mange mites, fleas of to Baker (1983), but on islands either in Lake many species, lice, small ticks, and viruses Michigan (Long and Long, 1986) or urban which may cause tumors. A disease called squir- islands in large cities it is not uncommon to rel pox was reported in Michigan (Stuht and see overpopulations of grays. Harte, 1973). Botflies occasionally leave their Gray squirrels are famous in mammalo- warbles in gray squirrels. Internal parasites in- gy as examples of mammals where numbers clude blood protozoans, roundworms, microfi- build up and the population, at least in part, laria, flukes, tapeworms, and spiny-headed moves away from one area to another, even worms (Kilham, 1959; Jackson, 1961; Scharf crossing rivers to escape population pressure. and Stewart, 1980; Clark, 1959; Parker and In early American history these movements Holiman, 1971; Koprowski 1949a). were often documented in the eastern United Home Range and Density. In Kansas, the States. In Michigan, Seton (1920) quotes from home range for gray squirrels was calculated the Bay City Tribune of February 17, 1907, as circular areas about 1.5/ha for females, that a population was observed in 1866, along and 3/ha for males (Armitage and Harris, a two-mile stretch of road to number 1,400 1982). Riege (1991) found that home ranges squirrels. Hundreds of black phase grays and overlapped considerably. The average mini- normal grays crossed the Raisin River (Wood, mal-area polygon for 11 squirrels was 1.5±0.8 1922). Banfield (1974) mentioned a 19th hectares. In my experience some squirrels stay century crossing of the Niagra River, and Fryx- on my acreage (2 ha) all winter, fattening ell (1926) mentioned grays crossing the Mis- themselves at my bird feeders; but some squir- sissippi into Iowa in 1905 and 1925. Jack- rels come and go, being observed as far away son (1921) mentioned squirrels crossing into as half a mile, and on rare occasions return- southern Minnesota. Many squirrels drowned ing to our property. Koprowski (1994a) men- in these movements. He also mentioned a tioned home ranges as 0.5 ha to somewhat horde of grays encountering Lake Chetek,

TAXONOMIC ACCOUNTS / ORDER RODENTIA 199 Barron County, while on the march, and a the taxonomy of the gray squirrel in Wiscon- dozen squirrels were observed swimming (one sin. The large subspecies in Minnesota, if it is at a time). About 20 were found drowned in a valid subspecies, is S. c. hypophaeus, de- bays and inlets. This emigration occurred in scribed by C. Hart Merriam. Wisconsin and late September and ended before October. the state of Illinois have been ascribed to two Long and Long (1986) reported a build up of separate races by Jackson and Hoffmeister, squirrels on Washington Island, resulting in an respectively. On what basis should one rec- exodus of grays swimming off into Lake Mich- ognize hypophaeus, of Minnesota, as a valid igan. Some (about a dozen) were observed in race, and where should the boundary between Ellersen Bay and others seemed to leave the it and the southern S. c. carolinensis be east side, more than 90 washing up on the drawn? Hall and Kelson (1959) and Hall beach of Rock Island (where no gray squirrels (1981) followed Swanson in restricting hy- occur). Apparently several hundred grays par- pophaeus to northern Minnesota, with its ticipated in this exodus. Shorger (1949) stud- eastward border the St. Croix River. This ar- ied the historical literature finding that gray em- rangement left both Wisconsin and Illinois to igrations resulted in the early 19th century S. c. pennsylvanicus. about every 1-5 years. There are no such em- Some have stated that the northern S. igrations occurring in recent years, except the c. hypophaeus (including in this concept Min- Washington Island irruption. Irruptions may be nesota, Manitoba, Wisconsin, and Upper more common on islands where populations Michigan) has a high frequency of melanistic fluctuate anyhow. The population numbers and squirrels. Speculatively, such a high number their crashes seem related chiefly to the avail- of black individuals could be attributed to the ability of mast as food for the squirrels. charred tree stumps and logs of northern for- Remarks. Dominance in fighting and ests after forest fires, or the ordinary expla- chasing does not seem to depend much on nation of Gloger’s Rule, that dark forms in- color in the gray squirrel. Both mutant yellow habit shadows of forests and well-watered and melanistic color phases described above regions (where the wet soils are dark). Black have been observed in some years to run away fur absorbs heat, not insignificant in squirrels from normal grays, and in other years either enduring the winters of the North Woods. In a yellow or black squirrel was observed to any case, melanistic squirrels are seen locally chase the grays. Oddly, the little red squirrel throughout the state of Wisconsin, extending Tamiasciurus even half grown will put a gray southward into Illinois in relatively high fre- to flight. I hypothesize that this may be be- quencies (Hoffmeister, 1989) and throughout cause of their color, which resembles that of the Upper Peninsula of Michigan. the red fox. Sciurus, although generally aso- Hoffmeister (1989) found the highest Il- cial, sometimes nest communally, especially linois frequencies of melanistic squirrels in Jo in winter. Koprowski (1996) noted “kin clus- Daviess, Carroll, Cook, Lake, and Adams ters” in gray squirrel mothers with adult young, counties, where most of these are found in but no fox squirrels remained in their “natal the northeast and northwest corners near areas” (philopatric). Unrelated adults nested Minnesota and Wisconsin. All the squirrels together in both species, but adult female fox shot in 1857, near the Rock River, about 50 squirrels rarely nested together. Related gray in number, were black (Kennicott). That is squirrel females showed no aggression toward (was) a significant frequency, unless they were one another; there was observed aggression shot selectively. Hoffmeister reports that to- among unrelated females. day the highest frequencies are along Lake Geographic Variation. The Jackson- Michigan from Evanston to Zion in Illinois. Hoffmeister pattern (Table Plan-1) is seen in Schorger (1949) believed the blacks were rare

200 THE WILD MAMMALS OF WISCONSIN south of Reedsburg and Baraboo in his day, size from north to south. I found none in Wis- and uncommon northward except where lo- consin (see Table Rod-5). cally abundant. Such places are Waupaca and Jackson (1961) studied grays in Wiscon- the zoo area in Wisconsin Rapids. I usually sin and Minnesota, and looked for other, more have one or two on my property in Stevens important (than melanism) characters. He Point, although for the first ten years there mentioned the dark venters as more common was none. I suspect in Wisconsin less than 5 in northwest Wisconsin rather in the south- % are black, except in some local populations. east part. He reported “distinctive color dif- Another northern characteristic is the ferences” in both winter and summer pelag- presence of conspicuous but hitherto unnoticed es. Not appreciated or overlooked were the winter ear tufts. Although not so large as in generally darker upper parts of hypophaeus; tassel-eared squirrels, they are indeed promi- the darker tail, particularly noticeable in the nent. Contrasting white and ochraceous hairs shorter white tips to the individual hairs and suggest some visual communication, and per- thus producing a much less frosted tail; the haps the tufts are used in species recognition base of tail hairs much more grayish and less and courtship, since tufts begin in late winter tawny or cinnamon; the brown tinge on the and occasionally last occasionally until early flanks darker and less ochraceous, the cheeks, June. The function of warming of the thin and temporal region, and ears brownish gray, of- prominent ears is obvious. I have never seen ten in winter distinctly iron gray, never yel- the tufts when I lived in Champaign-Urbana, lowish ferruginous or ochraceous as in S. c in central Illinois (three years), nor even when pennsylvanicus. I could not confirm any of I was looking for them in the winter of 1991, these differences as significant. at St. Olaf College, in southeastern Minnesota Havera and Nixon (1978) studied the (where I was Visiting Professor). On Washing- cranial dimensions (size) of squirrel skulls from ton Island, where some or all the grays may central Wisconsin, northwest Illinois, north have been introduced, most winter squirrels central Illinois and in the Illinois River Valley have tufts. In the southwestern Wisconsin coun- in northeast Illinois, extending a fair distance ties and in southern counties, in some popula- southward. They also compared with discrim- tions at least, no tufts were seen. This Muse- inant function analysis gray squirrels from the um’s collection contains some squirrels from remainder of Illinois with these northern pop- southern Wisconsin having tufts. Tufted squir- ulations. They noted a break in a cline be- rels were observed from Kenosha Co.; Janes- tween the northern populations and the Illi- ville, Rock Co.; Monroe Co.; Vernon Co.; nois River squirrels. These were intermediate Marquette Co.; Dodge Co.; Jefferson Co.; and between the northern squirrels and those Waukesha Co. No tufts were seen in winter southward. They concluded the northern pop- on specimens from Richland Co. ( 2 ); Dane ulations were in a northern race (which they Co.; Washburn Co.; Columbia Co.; and Mil- called pennsylvanicus instead of hypophae- waukee Co. A border specimen from Illinois us) and that the southern squirrels of smaller had tufts (McHenry Co.) but one from Jo size were referable to S. c. carolinensis. Daviess Co. (Stockton) did not. Hoffmeister (1989) rejected this classification. Hoffmeister (1989) studied size in gray The race S. c. carolinensis has extremely squirrels, for S. c. hypophaeus is supposed small individuals, and the dorsum is more or to be large, and he assigned all Illinois squir- less washed with tawny or tan brown. Any rels to the southern race S. c. pennsylvani- assignment of Illinois squirrels to this race was cus (although his map mistakenly refers all obviously incorrect. Assignment of most Illi- squirrels in Illinois to the race S. c. carolin- nois squirrels to S. c. pennsylvanicus is rea- ensis). He reported slight clinal variation in sonable. An assignment of the squirrels in

TAXONOMIC ACCOUNTS / ORDER RODENTIA 201 Table Rod-5Rod-5. Cranial measurements of some gray squirrels in Wisconsin and northern Illinois. There is no significant geographic variation evident in these and several other measurement. Sexes are combined. Generally the localities are north to south. Mean ± standard deviation and observed range. Illinois data are from Hoffmeister, 1989.

Locality N Greatest Zygomatic Cranial Int. L. Max. P4-P4 Postorbital (Counties) L. Br. Br. Br. Nasals toothrow Br. Br. Lincoln & 3 63.4 35.2 25.3 19.5 21.1 11.3 13.4 19.4 Marathon 62.3-64 34.9-35.5 25.8-26.6 18.2-22.6 18.8-22.6 11-11.7 13.3-13.5 18.9-19.8

Portage 12 63.1±1.49 34.8±0.7 25.9±2.15 19.51±1.04 21.04±0.55 11.52±0.35 13.18±0.45 19.57±0.9 59.9-64.6 33.6-36.2 19.3-27.2 17.3-21.2 20.1-21.7 11.1-12.03 12.5-13.9 17.9-21.2

Waupaca & 10 63.2±0.94 34.92±0.92 25.7±0.68 19.4±0.85 20.3±0.50 11.65±0.50 13.28±0.10 19.3±0.63 Waushara 62.2-65.1 33.7-35.9 25.6-27.4 18.1-21.0 19.5-21.4 11-12.3 13-13.7 18-20.7

Sauk 4 63.6±1.6 35.6±0.61 29.95±0.4 19.93±1.0 21.25±0.24 11.08±0.33 13.35±0.41 19.8±0.63 62-65.9 35-36.1 26.9-27.4 18.6-21.0 21-21.5 10.7-11.4 12.9-13.7 19.3-19.9

Marquette 3 62.7 34.5 26.1 20.3 20.3 11.0 13.2 19.8 61.5-63.7 33.5-35.2 25.1-26.8 19.8-20.9 19.6-21.8 10.9-11.1 12.9-13.5 19.2-20.1

Racine, 6 61.6-±0.8 31.46±0.69 25.38±0.33 18.73±1.6 20.2±1.16 11.36±0.49 13.13±0.54 19.25±0.56 Milwaukee & 60.6-62.6 33.8-35.6 25.1-25.9 16.7-21.3 18.7-21.0 10.7-12.1 12.5-14.0 18.7-20.2 Kenosha

Richland & 2 63.7 35.1 26.2 18.8 20.6 10.9 13.4 19.4 Vernon 62.6-65.8 33.4-36.8 25.3-28.0 17.8-19.8 19.9-21.3 10.8-11.0 12.9-13.8 19.2-19.6

McHenry Co., 1 61.5 35.0 26.5 18.2 19.4 11.2 13.2 18.8 Illinois

NW Illinois 18-22 62.4 35.0 24.4* – 20.3 11.3 – 19.4 * A low value. northern Illinois counties to S. c. hypophae- The traits of the northern gray squirrel us might have been made by Havera and Nix- populations are interesting microgeographic on. Cory (1912) showed little regard for hy- variation, but not constant enough for sub- pophaeus, saying it might be on average a specific recognition. little larger than typical squirrels from Wis- Specimens examined. Total, 112. Ad- consin. He ascribed the Wisconsin and north- ams, Ashland, Burnett, Chippewa, Clark, ern Illinois squirrels to a northern race (S. c. Columbia, Dane, Dodge, Door, Forest, leucotis (Gapper)). I cannot separate Wiscon- Green, Jackson, Jefferson, Juneau, sin squirrels from Illinois populations in size Kenosha, Lincoln, Marathon, Marquette, (Table Rod-5) or other characters. One should Menominee, Monroe, Oneida, Outagamie, understand that the discriminant function anal- Ozaukee, Pepin, Portage, Price, Racine, ysis, used by Havera and Nixon, which did Rock, Sawyer, Sauk, Sheboygan, Trem- separate northern populations of Illinois from pealeau, Vernon, Vilas, Washburn, Wash- central and southern samples, will combine ington, Waupaca, Waushara, Waukesha, and weight characters in such a way that a Wood counties. distinction can be made. Illinois and Wiscon- Illinois. Jo Daviess Co.: Stockton 1. sin squirrels are similar. McHenry Co.: No specific locality 1.

202 THE WILD MAMMALS OF WISCONSIN Sciurus niger Linnaeus The name Sciurus means shade-tail, re- Fox Squirrel ferring to its size and bushy quality, and the word niger means black. Fox squirrels are sel- Sciurus niger rufiventer Geoffroy dom black. The subspecific name rufiventer, St.-Hilaire means reddish belly, an apt description, except the color tends more to orange than red. 1803. Sciurus rufiventer E. Geoffroy St.-Hilaire. As in other Sciurus, there is a small nail Catalogue des mammiferes du Museum Nation- on the pollex, the third and fourth digits are al d’Histoire Naturelle, Paris. Page 176. Type elongate to encompass part of the circumfer- locality, Mississippi Valley. ence of a tree, and the eyes are large (giving 1851. Sciurus sayii Audubon and Bachman. The vi- a cute appearance and an alert expression). viparous quadrupeds of North America. 2: plate The bow-legged stance, adaptive for climb- 89 and page274. Type from river bottomlands ing, leads to the term scampering for running of Wabash River in Illinois, or from the Mis- about on the ground, which this species does souri River, or possibly Michigan. more than S. carolinensis. 1852. Sciurus Vulpinus Lapham. A systematic cat- The skull closely resembles that of the alogue..., Fourth Rept. Regents, p. 44. gray squirrel, but averages larger. The infraor- 1902. Sciurus ludovicianus Snyder. A list, with brief bital canal is smaller (less than 5 mm length), notes, of the mammals of Dodge Co., Bull. and the upper premolar 3 is absent. The cusps Wisc. Nat. Hist. Soc., 2:118. and cuspids of the molars are more rounded 1907. Sciurus niger: Osgood. Proc. Biol. Soc. Wash- or bunodont than the flat-worn cheek teeth ington, 20: 44. of gray squirrels. The chromosomes number 1908. Sciurus niger rufiventer: Jackson. A prelimi- is 40 (Nadler and Sutton, 1967). The bacu- nary list of Wisconsin mammals. Bull. Wisc. Nat. lum resembles that of S. carolinensis. The Hist. Soc., 6: 16. bones of the skeleton usually show a reddish- orange or “pinkish” tint (Levin and Flyger, 1971; Flyger and Levin, 1977). Does the pink in the bone relate to the red color of the blood? Haeme production, the source of the red, depends on the action of uroporphyringogen UI, chiefly in erythropoietic (i.e., blood-forming) tissues in bone marrow. Too much of UI results from insufficient enzyme uroporphy- rinogen III co-synthetase (or enzyme-UIII). Apparently the chemistry and titres in the fox squirrel resemble those in congenitally sick cattle and humans, who suffer “erythropoietic porphyria.” Overproduction of UI apparently is normal in fox squirrels, but some Wis- consin fox squirrel bones are not pinkish. The dorsal color of the pelage is ochracaeous gray, with a strong intermixture of black dorsally, concentrated on the forehead above the cheeks and eyes, and medially in the long tail. The toenails are conspicuously Sketch of fox squirrel Sciurus niger. By Francis L. Jaques. black against the bright orange feet, and the In Cahalane, 1961. vibrissae are long and black. The cheeks are

TAXONOMIC ACCOUNTS / ORDER RODENTIA 203 more or less orange, often dusky with suf- mae (Flyger and Gates, 1982). Measurements fused black, and the prominent ear tufts grown are in Table Rod-6. in winter and spring are bright orange. The Dental Formula. Upper third premolars venter is a rich ochraceous orange, paling to absent. I 1/1, C 0/0, P 1/1, M 3/3 = 20. rose-orange, almost pinkish orange in some Geographic Distribution. The distribution specimens. The tail is bright orange, almost of Sciurus niger, a tree squirrel preferring tawny orange intermixed with black or brown, forest edge habitats, is dynamic, as ever which is more visible dorsally than on the un- changing as the clearing of woodlands and derside. Distally the tail has long hairs tipped the growth of hardwoods. Additionally, there with pale ochraceous beyond the bands of is the possible displacement by competition black-orange-black hairs. Melanistic squirrels with Tamias and S. carolinensis, perhaps have been reported, but usually the black squir- some competition with other woodland mam- rels in Wisconsin are gray squirrels. A mutant mals (deer mice, flying squirrels, etc.). For pattern observed in Dane County, southwest example, at my home in Portage County, Wisconsin (La Crosse, also 2 mi. W Viola No. basically an oak-jack pine savanna habitat 4160), nearby Iowa (1 mi. SW Rossville), and overlooking a lake, gray squirrels are predom- reported by Kennicott (1857) in northern Illi- inant among eight squirrel species present, nois has black beginning on the cheeks and but only one fox squirrel has visited the place extending on the entire venter, and onto the in 23 years (staying only a day). Fox squirrels lower legs. The Iowan specimen has black on are found here and there in Portage County, the cheeks, intermixed on the crown a little, but gray squirrels are the abundant tree squir- and extends even partially onto the feet. The rels in deciduous woodland habitats or park face and body of this pattern with orange set areas. The only occurrence for this savanna against dark black is striking to say the least. area that I have observed in Portage County, It might function in display, in intimidation. was in woodlots or hedges adjacent to culti- Molt lasts about a month, beginning in vated fields, mostly toward Junction City and spring. New hair appears on the head and westward, and tamarack-cedar swamps east often a distinct molt line can be seen belting of Stevens Point. the body. Even young of the year molt and Cleland (1966) reported fox squirrels in grow a dense winter coat (with ear tufts). Some archaeological sites in a broad region from squirrels molt twice or late (e.g., Sept. 28, Ontario to Michigan, but Jackson (1961) found UW-SP 7131, Vernon Co.)(also see Koprows- the species confined to southern counties of ki, 1994b). There are eight blackish mam- Wisconsin before the turn of the 20th century. In oak and hickory woods they were abundant, occurring along the Mississippi to the St. Croix, extending into Dunn County, ranging southward as far as southern Juneau County, and along Lake Michigan as far as the Door Peninsula (even northward of Sturgeon Bay). Up to 1961, kill records were highest in this same oak-hickory and oak savanna region, but squirrels were killed even in northern counties (in low frequencies). No sign of them was in Iron, Vilas, and Florence counties. The observed expansion of range northward since 1961, was thought to result from Skull of Sciurus niger. cutting into the continuous northern forests,

204 THE WILD MAMMALS OF WISCONSIN formerly inhabited only by grays and reds. Coincidentally, fox squirrels swept northward in Lower Michigan, fastest along the lake shore counties, and by “introductions” colonized the Upper Peninsula in the easternmost counties (Baker, 1983). Fox squirrels reached the Up- per Peninsula by way of Wisconsin after publication of Burt’s work (in 1948). Baker (1983) reported that most of the forested Upper Pen- insula is “still not fox squirrel range.” Jackson (1961) reported a specimen from Brown Coun- ty near Green Bay, approximately 40 miles (= 64.4 km) from the Michigan boundary. Long

Maps showing distribution of Sciurus niger in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 205 (1974) reported them from both Marinette and Habitat. In a word, the fox squirrel pre- Florence counties extending the range to the fers “forest-edge.” Therefore, it is a mammal Michigan border. In Michigan, in the early of savanna, preferring open woods where it 1970’s, fox squirrels were noticed in the south- scampers about on the ground. A large for- western quarter of Iron County and southwest- est certainly is not required, and a hedgerow ern corner of Dickinson County. In 1977, a or fencerow with a few mature oaks is quite fox squirrel was found dead on a road in Bara- suitable. As one might expect, fox squirrels ga County, and another was found later in Me- also dwell in oak thickets, deciduous forested nominee County (Baker, 1983). wetlands with sunny openings, and other open Status. Fox squirrels have delicious flesh, deciduous woods. The dry, forest uplands of and were eaten in greater numbers than grays southwest Wisconsin, and the deciduous by pioneers (Schorger, 1949b). Today it is a woods in southeastern Wisconsin, where oaks, game mammal throughout Wisconsin and lo- walnuts and hickories are found, are favorite cally in the Upper Peninsula. Sport hunting is haunts of the fox squirrel. Where gray and its chief economic value. In natural woodlands fox squirrels occur together, fox squirrels may the fox squirrel is not so secretive as the gray, chase grays away, but usually the grays de- and forages later in the day. In the southern fend the nest cavities in trees, constructing counties where hunters seek bobwhite, par- relatively few leafy nests, and the fox squir- tridge, and especially the wild turkey (in oak- rels must utilize leafy nests in the sunny tree hickory and bass-maple forests near forest canopies. This sympatry is common in the edge, on sunny, rocky hillsides, and in cleared woodlands of west-central and southwest Wis- and disturbed habitats), many fox squirrels are consin. Hazard (1982) reported neither spe- shot as game. Some hunters seek them out cies dominates the other by fighting and chas- specifically, sitting in the woods where tree ing. Steele and Weigl (1992) discussed ener- nests are conspicuous, waiting until one scam- getics and patch use of the habitat. In wood- pers out in the sunshine, and shooting them lands, especially continuous hardwoods, the with rifles equipped with scopes. In most coun- grays usually win out in Wisconsin. In fir-ce- ties the rarer fox squirrel is taken less frequently dar and pines the reds usually win out. Either than grays. The fur is of little value, and is not or both Sciurus may thrive in a town, but saved in Wisconsin. Once the tails decorated usually in Wisconsin the grays are established car antennae, before the advent of Green Bay in urban areas. Packer-mania and the use of Packer flags. The One may evaluate habitat suitability in fox squirrel causes some economic damage by the fox squirrel by reference to canopy of gnawing to human habitations, and some fruit trees, including those with food sources, es- trees are ravaged. In corn-growing areas, which pecially oaks, and to nearby cultivated grain seemingly encourage fox squirrels to move into crops. These plants provide essential winter adjacent fencerows and wood lots, fox squir- foods. Understory must be present for cover rels may eat enough corn to cause significant and reproduction. Fox squirrels’ nesting hab- damage. Aesthetically, the fox squirrel is a its require fewer nest cavities, but the fox squir- beautiful mammal with its orange ear tufts rel will use them. Closure of canopy may range prominent in winter, orange coloration and from approximately 22-60 %. Shrubby un- long, lovely tail. Few can fail to appreciate its derstory is of little use to this species if it ex- beauty either as it gracefully climbs about in a ceeds 30 percent. tree or sits erect, neck outstretched, standing Tree nests are quite often constructed in on its hind limbs on the ground. Squirrels are branches of trees about 30 feet or higher above second only to songbirds as objects of photog- ground, and are made of shredded leaves, bark raphy (Koprowski, 1994b). cut from twigs, and branches. Hoffmeister

206 THE WILD MAMMALS OF WISCONSIN (1989) states that in the tree cavity the nest of ki, 1994b). The young are born in nest cavi- a fox squirrel is usually made of leaves. ties of trees and occasionally in tree nests Foods. Foods in Wisconsin are similar to (Allen, 1942). The newborn young weigh those of grays, including primarily the mast about 15 grams, and are blind and naked. In of the hardwood forests. Acorns, maple seeds, a week hair appears on the back, and the buds of elm and oaks, cambium bark and sap weight doubles. By 35 days the young are (like the gray squirrels, the fox squirrels often active, eyes open, and the body hair is almost gnaw on the cambium layer of trees and eat furred out. Allen (1943) mentions that the eyes sap in spring and winter), mushrooms, arthro- open by the fifth week. By 55 days the young pods, and such foods are often eaten. Fruit emerge from the nest, about 100 days after from trees, corn and many kinds of nuts and conception. Weaning follows three months seeds are also eaten. Allen (1943) lists im- after birth. Pregnant females drive away their portant mast bearing trees for Michigan, young, but second litters may accompany the which also occur in Wisconsin: white, red, bur mother several weeks longer than usual. An- and black oaks. nual production is about 6 young per adult Allen (1943) and Reichard (1976) men- female, less in yearling females, which com- tion such diverse foods as flower buds of ma- prise about 60 per cent of females in Michi- ples and bur oaks, maple sap, catkins of wil- gan (Allen, 1943). Fox squirrels survive more low, cottonwood, flowers of beech, elm, hack- than 6 years (Fouch, 1958), up to 9 in cap- berry and basswood, beetles, tubers, bulbs, tivity (Crandall, 1964). roots, nest eggs and young of song birds, and Mortality. Few predators can catch tree seasonal strawberries, serviceberries, haws, squirrels regularly. Probably on rare occasions plums, raspberries, blackberries, greenbriar, a sick, old or very young individual is caught cherries, ash, dogwood, elderberries, and oth- by raptors (hawks, occasionally owls, once an ers. Acorns, nuts, seeds, and corn are also osprey) and Carnivores (especially foxes) (Jack- preferred fare. Fungi, grapes, wild rose, and son, 1961; Allen, 1943). Domestic dogs and bittersweet are eaten now and then. Many cats kill them, but the greatest mortality to- food items are listed by Koprowski (1994b). day is from automobiles and hunters. Reproduction. Fox squirrels have two lit- Jackson (1961) lists many parasites in- ters each year (Allen, 1943; Brown and Yea- cluding lice Enderleinellus and Neohae- ger, 1945; Koprowski 1994b). The first sea- matopinus, mites Euhaemogamasus and son of breeding begins in January and gesta- especially Sarcoptes scabei (for the mange), tion lasts 44 or 45 days following mating. The flea Orchopeas, chigger Trombicula, and second season begins in June and lasts some- ticks Haemolaelaps and Dermacentor. A times into October. A specimen from near variety of internal parasites include blood pro- Nichols, in Outagamie County, was lactating tozoans, spiny headed worm, 10 species of on October 28. Yearling females usually breed round worms (including Bohmiella, Entero- only once. Breeding is diminished by low food bias, Helig-modendrium, Physaloptera, and supplies and bad weather. Breeding is en- Trichostrongylus that occur in Wisconsin hanced by previous removal of adults, espe- squirrels). Sometimes the incisors do not oc- cially females (Hansen and Nixon, 1985), clude, and grow in curls, even penetrating the which are the most stable and territorial of squirrel’s brain. Koprowski (1994b) lists pred- the population. Litter size varies up to 7 (one ators, parasites, viruses, and pesticide prob- specimen had 7 fetuses, Hoover, 1954). Av- lems in great detail. They often work in com- erages have been reported between 3 and 4 bination to kill off populations. in central Illinois (Hoffmeister, 1989), but over Allen (1943) discussed how winter and its range the mode is usually 2 or 3 (Koprows- food scarcity related to predation and scabies

TAXONOMIC ACCOUNTS / ORDER RODENTIA 207 on weakened squirrels would increasingly af- squirrels. Koprowski (1994b) mentions home fect fox squirrels, which are southern mam- ranges for this species from 0.85-17.2 ha (2- mals dispersing northward into the North 40 acres) for females and from 0.54 to 42.8 Woods. A severe winter (1935-6) was related ha (1-100 acres) for males. C. E. Adams to a decline in fox squirrels in Michigan. Sca- (1976) presented some characteristics of bies (sarcoptic mange), an obvious factor, fol- home range for the fox squirrel. Home rang- lowed a winter of low oak mast. es overlap extensively, and territoriality of Much more research is needed on the areas is not evident. Fighting over females is competitive interactions of several kinds of another kind of territoriality. Juveniles disperse squirrels and other mast eaters in Wisconsin. from home areas to new homes. Males dom- Periodic mast scarcity is in itself a great prob- inate females, and adults intimidate juveniles. lem for S. niger and other squirrels. Allen Remarks on Behavior. Hazard (1982) (1943) discusses this thoroughly, and some doubts that interspecific fighting in squirrels of his ideas are included here. “In 1939 there results in one species’ dominance. In the fox was the greatest abundance of mast occur- squirrel population, adult females are the most ring in five years, and in 1940 the population stable in their home ranges or territories (see probably reached a high for the same period. below). Either in courtship chases or territori- But the complete failure of the 1940 acorn al battles, or even in displacement during food crop precipitated a crisis in the ensuing win- competition, two adults begin with tail flick- ter. Among fox squirrels there were many ing, as in grays. Purrs, barks, and other scold- signs of malnutrition and weakness; 16 indi- ing sounds are emitted during courtship, per- viduals were found dead in natural dens, and formed with various approaches, probably wood duck and raccoon nest boxes. Five more scent emission, and even courtship groom- were found on the ground. In addition, a gray ing (McClosky and Shaw, 1977). Jackson squirrel and two flying squirrels were record- (1961) reports that the fox squirrel scold is ed as mortalities. Mange was wide-spread and more guttural and lower in pitch than that of those autopsied were 3 to 8 ounces (87-232 the gray, ending in a drawn out “qwa-a-a-a- g) under the average weight for late winter a.” Although the species is said to not be the year before.” Other details of starvation, strongly territorial (for example, Baker,1983) emaciation, and convulsions are described by the female den sites are evenly spaced (about Allen. Scabies and shock were considered 50 m apart). Nevertheless, the squirrels for- extremely dangerous to squirrels. In addition age in overlapping home ranges, and in win- to direct mortality, “food scarcity further re- ter several may sleep together in their nests. duced the population by preventing breeding.” The orange winter tufts in most Wiscon- Extreme winter weather works adversely with sin fox squirrels, for one thing, are for warmth, food scarcity against fox squirrels. and even the pinnae of the ears may be furred Allen (1943) mentions the weakness of (Flyger and Gates, 1982). As in the gray squir- emaciated squirrels caused by mast failure, and rel, S. carolinensis, the prominent tufts, additionally the ice and snowcover that buried which are even longer than in the gray squir- nuts so deeply they are “less available.” The rel, may be related to some overt or positive general weakening also makes squirrels more visual communication. susceptible to predation by dogs, hawks, or owls. My brother David Long and I raised Kan- Home range and Density. Jackson san fox squirrels for pets. One that escaped (1961) gives home range as about 3 or 4 from him bit my mother in the face. Another acres, occasionally exceeding 20 acres (80 pet fox squirrel bit Hartley Jackson (1961) in ha). Wise (1986) reports that fox squirrels the face. A person should not expect docility wander in larger home ranges than do gray from an adult fox squirrel. But one I had was

208 THE WILD MAMMALS OF WISCONSIN Table Rod-6Rod-6. Some measurements of Sciurus niger from Genus Glaucomys Thomas Vernon and Manitowoc counties, Wisconsin (3 males). North American Flying Squirrels

TL Tail Hindfoot Ear Gr. L Zygomat. br These softly colored, pug-nosed but lovely 591 290 66 22 66.7 36.8 squirrels are characterized by a lateral fold of 702 265 75 26 — — loose skin, which stretches between wrist and 592 271 65 24 — — ankle. Some workers call it a “plagiopat- agium.” Resembling the calcar of the uropat- excellent, in the University of Kansas gradu- agium of bats, there is a process (Baker, 1983) ate student dorms where I lived. It would car- that helps to make the membrane taut. Dis- ry items my wife was ironing up to the top of covered by Muul (1968), as far as I can deter- the bedroom drapes. mine, the unnamed wrist structure might be Fox squirrels, consistent with their pref- called a carpar analogous to the bat’s calcar. erence for the forest edge, spend a great deal The presence of a flight membrane, used to of time foraging or scatter hoarding nuts on glide from high elevations, as in several other the ground. Reportedly they run from enemies mammalian groups, was considered sufficient- over the ground longer than grays, eventually ly divergent to arrange these squirrels in a taking to the trees to escape. For communal separate subfamily (Petauristinae). Because of nesting and philopatry, see Koprowski (1996). this membrane, the crepuscular or nocturnal Geographic Variation. In Wisconsin there habits instead of diurnal behavior, and by the is no significant geographic variation in the consequentially enlarged eyes and flattened fox squirrel, and none would be expected in tail, Long and Captain (1974) and Black populations descended from southern popu- (1963) proposed that the flying squirrels, lations dispersing northward in historic times. known from North America and Asia, were No significant sexual variation was noted. directly derived from tree squirrels (such as Specimens examined. Total, 61. Adams, Sciurus). Therefore, Glaucomys and the Ori- Brown, Burnett, Crawford, Dane, Dodge, ental gliders are not assigned to a separate Green, Green Lake, Jackson, Juneau, Ke- subfamily. waunee, La Crosse, Langlade, Manitowoc, The skull in Glaucomys is small with Marquette, Marathon, Menominee, Monroe, delicate postorbital processes and downturned Outagamie, Pierce, Portage, Richland, Sauk, Vernon, Vilas, Waupaca, Waushara, Washington, Winnebago, Wood counties.

Gliding Flying Squirrel. Walter Weber, in K. Schmidt. Glaucomys sabrinus. By Susan E. Smith. 1992.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 209 nasals. A tiny upper third premolar is present, arched skull of G. volans has a proportional- as in the tree squirrels. The fur, exceptionally ly larger orbit, due in part to the narrower soft and fleecy in the North American Glau- jugal which is developed less anteriorly than comys, is furred in such a way the hairs com- that in G. sabrinus. Superficially it would seem prise an air foil for gliding locomotion. from examining the skull that G. sabrinus would have more difficulty seeing forward. A conspicuous difference is in the color of the Glaucomys sabrinus (Shaw) tail, which dorsally is not brighter and red- Northern Flying Squirrel dish as is the dorsum, but dark, dull walnut brown. Occasionally the reddish of the dor- 1801. Sciurus sabrinus Shaw. General Zool., 2:157. sum extends onto the base of the tail, but not far. In comparison, the tail of G. volans is Glaucomys sabrinus macrotis (Mearns) brighter than its dorsum, a pale tawny gray. The underside color of the tail in volans is in 1898. Sciuropterus sabrinus macrotis Mearns. Proc. contrast to the soft, tawny gray color above, U. S. Nat. Mus., 21: 353. Type from Hunter instead a lovely ochraceous or gray-orange. Mtn., 3300 ft., Catskills, Greene Co., New York. In G. sabrinus the underside is brownish gray, 1915. G [laucomys]. s[abrinus. macrotis: A. H. ranging from almost black to a dirty white. Howell. Proc. Biol. Soc. Washington, 28: 111. The skull has a small, downturned rostrum, with the braincase twisted ventrally, so The name Glaucomys means gray that in lateral profile the rostrum and cranium mouse, and sabrinus may mean something seem flattened and short. The remainder of like Severn, which is the type locality (mouth the skull is massive. The postorbital processes of Severn River, Ontario). The namer Shaw are attenuate, flaring away from the cranium. did not say, but Baker (1983) suggests the The nasals and maxillary extensions are aligned old Latin name for an English River changed with the zygomatic plates, anterior to the zy- from the Roman to English Severn probably gomata. The zygoma (especially the jugal bone) indicates the Canadian Severn River. The race is relatively thick. These cranial characters use- name macrotis means large ear. fully distinguish the species from other sciu- Description. The northern, and also the rids, except not so much from G. volans. southern flying squirrels, are readily distin- The dorsal pelage in Glaucomys sabri- guished from other sciurids by the lateral nus is grayish overlain with cinnamon-ochra- membrane used to glide, by enormous eyes ceous color, even chestnut in bright, new for nocturnal vision, and by large mouselike pelage. The pelage is darker in fresh molt, ear pinnae. The northern flying squirrel is sig- where the cinnamon brown tips are not grown nificantly larger than the southern. The ven- out, and in worn pelage where the tips seem tral hairs are more grayish at their bases, so to be worn off. The brightest chestnut colors that the venter is more often mottled with are seen in July pelages, and in one speci- grayish, even on the throat. The baculum is men taken in early September. In Septem- short and slightly twisted, and relatively stouter ber-November specimens the fur is significant- than in G. volans. It measures only 6.3-7.3 ly darker plumbeous or blackish gray in tone, mm (Burt, 1960). There are four pairs of from the underlying dark color. The tail re- mammae in the female. sembles the dorsal fur above, but may be more The two species of flying squirrels are ochraceous. On the neck the ochraceous also proportionately similar, as has been often may be brighter. The cheeks may be grayish. noted. I submit two characters which seem to The ventral fur is a dirty white due to the gray distinguish them in Wisconsin. First, the small, bases of the hairs. The neck area is more

210 THE WILD MAMMALS OF WISCONSIN nearly whitish, but even here gray is evident. deed, the distribution of Glaucomys sabrinus The dorsal pelage is brighter and more red- is more likely receding and the numbers prob- dish than in the drab, soft-colored G. volans. ably diminishing. This problem is very possi- Additional remarks on color are given in the bly developing because (a) the related south- account of G. volans. An annual molt occurs ern flying squirrel, G. volans, is extending its in late summer and early fall. actual range northward into the range of G. Measurements and weights are given in sabrinus, and (b) there is direct competition Table Rod-7. The measurements of the skull between the two species, for example, nest- and other bones are significantly larger than ing cavities (tree cavities), usually taken over in G. volans, but proportionally there seems by the earlier breeding G. volans (Muul, no way to distinguish the two species (except 1968). In Central Wisconsin, there are evi- in the form of the baculum) from one another. dences that volans adapt better to the spread- Dental Formula. I 1/1, C 0/0, P 2/1, M ing human settlements. 3/3 = 22. The anteriormost premolar is The proper management and conserva- minute. tion of this remarkably lovely, interesting, and Geographic Distribution. The scarce harmless sciurid, then, promises to be a real northern flying squirrel is found in boreal wood- challenge. The northern flying squirrel must lands northward of a boundary extending from not only adapt to the activities of humankind, Burnett County southeastward into Portage but it is contested by a near relative, which County, possibly northern Juneau County, may eradicate this rarer, boreal species. Pos- thence into Outagamie County and probably sibly some nesting-house design may prove not continuously throughout Door County (ap- beneficial, and in remote and continuous parently restricted to the forests north of the woodlands bolster the fecundity of G. sabri- canal at Sturgeon Bay). Not found on any Wis- nus. The species should be placed on a list of consin islands in Lake Superior (which is sur- threatened mammals in Michigan and Wis- prising to me) or Lake Michigan (which is like- consin (Long, 1974). wise surprising, because the islands are close Judging by museum collections, in which together, the species is the boreal one, and never are there more than a few of these the squirrels do not hibernate). animals represented from any locality, the Status. The geographic range today is total number of animals preserved is much about as it was in the years preceding Jack- lower than is the number of southern flying son’s work. The slight extension of range squirrels (G. volans). In Portage County, southward is likely that of known range. In- where both species now are found sympatrically, G. volans is much more abundant. This may to some extent reflect the greater abundance of volans near man’s habitations, and doubtless reflects the localization of G. sabrinus in conifer communities and wet forests north of Stevens Point. There are estimates that the number of G. sabrinus has been reduced in Wisconsin (Jackson, 1961) and in Michigan (Muul, 1968) from earlier days. I have found no evidence to the contrary, and I do have evidence that the southern species is moving northward. Habitat. According to Jackson (1961) the Skull of Glaucomys sabrinus. habitat preferred in Wisconsin is mixed hard-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 211 wood and conifer, usually a wet forest, seldom a pure hardwood stand, usually forests where decayed logs lie on the forest floor. The trees are usually hemlock-maple or hemlock-gray birch, and occasionally cedars. In Michigan, this squirrel has been reported in conifer swamps (Green, 1925) and in jack- pines, cutover hardwoods, and mixed conifer-hardwood forest (Manville, 1948). There are two kinds of homes. The cavity is used almost any time of the year, in live or dead trees, hollow stumps, the tops of snags or telephone poles, and nooks in the attics of

Maps showing geographic distribution of Glaucomys sabrinus in Wisconsin and North America.

212 THE WILD MAMMALS OF WISCONSIN old houses. The cavities may be natural, or parently breeding re-commences in summer. remodeled woodpecker holes. Nests are found Gestation lasts about 37 days according to Muul in the tree cavities, for newborn litters. The (1969). Specimens taken from a litter of six, stick nest (comprised of twigs, lichens, moss- from Woman Lake, Cass Co., Minnesota, were es, and shredded bark, Howell, 1918; Cow- tiny on May 20, 1950. Respectively, Univ. an, 1936) is made in spring and summer, Minn. 2901, 2900, had these standard mea- usually in evergreens high above the ground. surements and weights: 151, 151; 64, 63; 26, The nest of another squirrel or a crow may 25; 26 g, 26.2 g. After three weeks in captiv- be used. An opening facing the trunk is usu- ity at reportedly 43 days of age, a third specially made for rapid exit in the dreys. In win- men was 183, 82, 28, 32.7 g. ter, up to nine squirrels have been found in Newborn young are about 70 mm in one tree cavity (Jackson, 1961). length, weighing about 5.8 g. They are na- Foods. As in other tree squirrels it is very ked with eyes and ears closed, but the pata- important for G. sabrinus to cache seeds, co- gial membrane is evident (Muul, 1969). In six nifers seeds (usually in the cones), and acorns days vibrissae are evident, and in 18 days the in large quantities for lasting the winter. Foods pink bodies are furred. The eyes open in 31 are actively procured even on cold nights, tak- days, and the flattened tail is prominent. Af- en from boreal trees (such as spruce seeds) and ter 40 days the young emerge from the nest, supplemented by buds, arthropods, especially and weaning seems complete by 60 days. The moths and beetles, and dripping maple sap in young glide about after about 90 days, and season. Bird eggs and young are reportedly leave the parents at this time. Squirrels in this eaten, and certain edible fungi are often eaten collection were most numerous in October, when available. Where beech trees are found including young-of-the-year. the seeds are stored for winter in hollow trees Mortality. The barred and great gray owls and in stumps. Occasionally pine cones are (Seton, 1953; Corace et al., 2007), gray wolf piled on the ground. If G. sabrinus finds the (Voight et al., 1976), bobcat (Pollack, 1951), cache of another species of squirrel it raids it and domestic cats (Toner, 1956) prey on G. (Brink and Dean, 1966; Smith and Aldous, sabrinus. Large fish may occasionally eat one 1947; Wrigley, 1969; and Seton, 1953). that falls into the water (Seton, 1953; see Reproduction. G. sabrinus usually has a Baker, 1983). Other owls and nocturnal fur- single litter in late spring, but some females bearers doubtless take them, and many pred- bring forth a second litter in late summer. If ators conceivably eat the young. A forest fire there is a second, perhaps the first was lost might kill these squirrels. Rarely while gliding (Banfield, 1974, Doutt et al., 1966). The spec- some are impaled by sharp objects. Fleas imens in this University’s collection show no (Manville, 1949), lice, mites, chiggers, tape- records of pregnancy, and few adult records worms and roundworms may parasitize these before mid-July. Lactating females have eight squirrels. Inasmuch as the northern and south- teats, suggesting about four embryos more ern flying squirrels occasionally are sympatric or less. Muul (1969) reports the number is in northern and central Wisconsin, and the about 3 (2-6). Lactating females were ob- kinds of parasites are so similar, cross-infec- served 14 May, 2 September, and 13 Octo- tivity is a possible danger to one or the other. ber (which date is surprisingly late for such a J. N. Pauli, S. Dubay, E. Anderson, and S. J. boreal mammal). Courtship begins in March, Taft (personal communication) appraised the probably even before the thaws set in, and incidence of the parasite Strongyloides ro- continues until late May. The male stomps its bustus in fecal samples of live-trapped Glau- feet and emits high, pulsed calls. During cop- comys volans and G. sabrinus in an area of ulation females purr and males whine. Ap- sympatry (Schmeekle Reserve, Univ. Wiscon-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 213 sin—Stevens Point) hoping to document a lade, Lincoln, Oconto, Oneida, Portage, detrimental effect on sabrinus that might limit Rusk counties. its distribution southward. However, about Michigan Records. Schoolcraft Co., 4 half of G. volans was infected, whereas only mi. N Manistique. Houghton Co. (Crider, 11 percent of sabrinus was. This work may 1979, unpublished study on the Sturgeon be published more detail and fails to support River). Marquette Co. (Haveman and Robin- the hypothesis that one squirrel replaces the son, 1976). Menominee Co.: (Burt, 1946). other with the help of the parasite [as both See also Stormer and Sloan (1976). squirrels have been sympatric in this area for Other records. See Jackson, 1961; Mc- many years]. Cabe 1981: Forest Co., Sea Lion Lake; Flo- Home Range and Density. Hardly any- rence Co. Lost Lake. Sight records, question- thing is known of the northern flying squir- able, of squirrels reported to me from Juneau rel’s wanderings, in Wisconsin. In far off North and Monroe counties. Carolina and Pennsylvania, where home range was determined by radio tracking, it was estimated to vary from 8 to 32 acres (3.2 / Glaucomys volans (Linnaeus) 12.8 ha). Manville (1944) estimated that G. Southern Flying Squirrel sabrinus numbered 1/14 acres (1 / 5.6 ha) on two study areas in the Huron Mountains. Glaucomys volans volans (Linnaeus) Remarks. The flying squirrels are nocturnal and glide or voloplane dextrously from 1758. [Mus] volans Linnaeus. Syst. naturae, 10th ed. high to lower elevations as a means of loco- 1: 63. Type locality fixed by Elliot (Field motion, besides climbing with agility in trees. Columb. Mus. Zool. Ser., 2: 109: 1901) in The northern flying squirrel spends more time Virginia. on the ground than the southern. 1915. [Glaucomys] volans: A. H. Howell. Proc. Biol. Additional Natural History. Wells-Gosling Soc. Washington, 28: 109. and Heany (1984) reviewed the natural history of Glaucomys sabrinus. The name Glaucomys means gray Geographic Variation. None was detect- mouse; volans means to fly. ed in Wisconsin. The northern flying squirrels Description. This small, pug-nosed, ar- in Wisconsin have been referred to the race boreal squirrel is characterized by a lateral fold G. s. macrotis, an eastern race with type from of skin covered with fur, with which the squir- New York, by Hartley Jackson (1961). A. H. rel glides as one means of locomotion. Closely Howell (1918) drew a boundary line between resembling G. sabrinus (above) especially in macrotis and the nominate subspecies in north- cranial proportions, G. volans is significantly west Wisconsin. There is no mention ever dis- smaller. The hind foot is less than 33 mm, cussed by Jackson as to where to draw that the skull less than 37 mm; and the maxillary boundary. The racial boundary probably lies toothrow is less than 7 mm. The skull is not northwest of Wisconsin. Jackson found no dif- only smaller, but the orbit relatively large ferences between northwestern specimens and owing to a slender jugal bone developed less specimens from Door County. The nominate at the anterior end. There are eight ventrally race is probably larger and darker in color of arranged mammae (in four pairs). There are pelage than seen in Wisconsin flying squirrels. 48 chromosomes, according to Hoffmeister Skull measurements for Wisconsin specimens (1989). The baculum is slender, and slightly are listed in Table Rod-7. twisted. Although G. volans is smaller than Specimens examined. Total, 24. Bay- G. sabrinus the baculum is longer (12.1-12.8 field, Clark, Door, Douglas, Forest, Lang- mm) (Burt 1960).

214 THE WILD MAMMALS OF WISCONSIN In Wisconsin, there are apart from size Geographic Range. At present the south- differences, significant differences in color. ern flying squirrel is to be expected in any See the account above of G. sabrinus, par- county in Wisconsin, and is eventually spread- ticularly the comments on the color of the ing its geographic range throughout much of venter and tail. Dorsally, the color in G. vol- the Upper Peninsula of Michigan. Previously ans is paler, more of a cinnamon pecan or known only in southern Wisconsin, the spe- pecan color, whereas in G. sabrinus the col- cies has been dispersing northward possibly or is richer chestnut or ochraceous and in at least in part at the expense of G. sabrinus. worn pelages darker, walnut brown. Ventral- See Map. So far the species is restricted from ly, the color of G. sabrinus is splotched gray- the Door Peninsula and some heavily forest- ish because the bases of the hairs are ed northern counties, and from all the islands plumbeous gray. In G. volans, the color is more investigated in Lake Michigan and Lake Su- whitish, seldom is there an intermixture of gray, perior. The record from Burt (1948) in south- and especially on the throat the hairs are pure eastern Upper Michigan suggests that former- white. The underside of the tail tends more to ly there was a connection between that pop- pinkish cinnamon, or ochraceous, less washed ulation and the Wisconsin southern flying with blackish. It is a lovely color scheme. Molt squirrels southward and southwestward, or seems to begin in late summer in the old adults, possibly with populations spreading along but there is little known about molting in G. Lake Michigan from Lower Michigan rang- volans, especially in Wisconsin. ing southwestward to the Wisconsin border. The southern flying squirrel is one of the Status. The southern flying squirrel seems smallest of the Wisconsin sciurids. The exter- abundant locally, especially in the southern nal meaurements are approximately 220-253, counties and near human habitations. The 80-120, 28-33, which are smaller than in G. species is spreading its range northward, in- sabrinus (260-360, 115-170, 34-45). Mea- vading the forest counties previously occupied surements are in Table Rod-7. only by G. sabrinus. No protection is deemed Dental Formula. As in G. sabrinus, the necessary, inasmuch as the common South- formula is as follows: I 1/1, C 0/0, P 2/1, M ern flying squirrel may be increasing at the 3/3 = 22. The anteriormost premolar is a expense of the northern. The species seems tiny spicule. harmless to man, although sometimes in winter raising our concerns by sleeping and gath- ering in people’s attics. There is some predation on birds in nestboxes. There is no eco-

Glaucomys volans in a bird box Roger Tory Peterson. Courtesy of Virginia Peterson, Houghton Mifflin. Skull of Glaucomys volans.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 215 nomic value for fur or food, but aesthetically flying squirrels are beautiful and fascinating. Habitats. A denizen of the hardwood forests, especially those in southern Wiscon- sin (hickory, maple, oak) the geographic range differs dramatically from G. sabrinus in being largely congruent with the prairies. This is true not only in southern and western Wis- consin, but also in the prairies along Lake Mich- igan extending into Michigan. This pattern seems odd because the squirrel is arboreal. Some adverse ecological factors that hypothetically may be involved with the prai-

Maps showing geographic distribution of Glaucomys volans in Wisconsin and North America.

216 THE WILD MAMMALS OF WISCONSIN rie distribution are the prior occupation of G. actually hibernating and are often active on sabrinus in the northern woodlands in the cold nights. Nests for bringing forth young, absence of G. volans; pine-maple instead of in birdhouses, woodpecker cavities, and nat- oak-maple forests in the north; recent inva- ural tree cavities, even in the dreys of other sions of prairie country in Wisconsin follow- squirrels (doubtless abandoned), are usually ing the riparian woods and deciduous forest made of leaves and shredded materials, such edges; and dispersal following planted trees as the inner bark of trees. One nest (see Fig.) of farms extending the range northward pri- found in a bird house and preserved in the marily in the cultivated prairie regions farm UW-SP collection is about 7 inches in diame- by farm. Mapping the geographic range from ter rolled up around the edge and depressed a collection of specimens documents the squir- in the middle, comprised of shredded inner rel primarily from populated areas (non-con- bark (May 7). Two others (in May and on Aug tinuous forest). In Illinois, second growth hard- 7) were made of the same material. A fourth woods are preferred, although not required found 26 May in a bird house abandoned by for this squirrel. Brush seems essential chickadees incorporated their moss nest with (Hoffmeister, 1989; and others). shredded bark added from above, which even- Both flying squirrels occur together in tually became well mixed at the bottom of the white pine-tamarack-maple woodlands and nest by the nestlings. A fifth was in a box marshes just north of Stevens Point. On the appropriated from chickadees (broken eggs), Schmeekle Nature Preserve; G. volans is for on July 25, a mother squirrel was found more abundant, and is distributed through- in the mossy nest with five newborn young. out most of Portage County as well. Young squirrels are often moved to new nests In winter, the southern flying squirrel may (Madden, 1974), but in every case I observed, be somewhat communal and is often found the mother is attentive when her nest is dis- in groups in bird houses, tree cavities, and turbed, never over 2-3 m away, and she al- attics. I have heard of numbers approaching ways returned to it. 30 for attics in Portage County and one house Foods. As in most arboreal squirrels, the somewhere in northern Wisconsin. Jackson chief food is acorns, supplemented with hick- (1961) reported as many as 22. During tor- ory nuts, maple seeds, fungi, berries, inverte- por the body temperature drops for short brates, and bird eggs and young. When food periods (Muul, 1968), but the squirrels are not is scarce buds and bark are eaten (Seton, 1953; Dolan and Carter, 1977; Banfield, 1974; Muul, 1968). The squirrels harvest and cache in systematic procedures (Avenoso, 1968; Muul, 1968). In late May, 1999, when my wife and I discovered the mother flying squirrel with young in a birdhouse formerly occupied by chickadees, and their mosses were incorporated with the squirrels’ shredded wood and bark, the question arose, was the house appropriated or abandoned by the birds? That same week there was predation in a nearby birdhouse occupied by tree swallows (Tachy- cineta bicolor) where six newly hatched young had been fed by the parents. Inside the box, Shredded wood nest of Glaucomys volans. one dead parent was found partially eaten.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 217 Six young lanquished and died, and there was the young. By May 1, the young were haired no sign of the other parent. The opening of out, eyes open, and the tiny ribbon like tails the box was so small that only a southern fly- were twined about one’s finger. They watched ing squirrel or a small weasel could enter, and us closely as we handled them. This handling the latter is not only rare in this area but would was repeated until 6 May. On May 7 the fam- have a difficult time ascending the 1"- wide ily had departed, after some three weeks un- metal pole. There were telltale gnawings at der observation. Ants had invaded the nest. the entrance. The putrid carcasses may have Another nest found by my wife and me, been the reason no squirrels nested in this on 25 May 1999, contained two young near- box. The next day, the nestbox housing an- ly as large as the mother, and they bobbed other family of black-capped chickadees their heads up through the shredded bark to (Parus atricapillus) was also emptied, with watch us, then ducked in again, while the some crushed eggshells present. The evidence mother leaped to the fence, about a meter suggests the southern flying squirrels de- away, watching us with her huge eyes, with stroyed two and perhaps three nesting fami- the sun glowing through her eight translucent, lies of birds. About three weeks later the box elongated, pink teats. Another nest found 25 contained a mother squirrel with five newborn July, contained slightly furred young, with young (see Homes above). In June, 2000, naked patagia resembling bats, about 55 mm another birdhouse was taken over from tree in length, and with tail long and nearly round. swallows with young, and the mother swal- One day later the tail seemed slightly flattened. low had been killed. The naked patagia extended forward of the Reproduction. Hardly any information is forelimbs onto the neck and posteriorly a few available from Wisconsin, and my specimens mm behind the thighs, approaching the tail. do not add much. My son John, his children, The eyes were shut. One young squeaked nois- my wife and I observed a litter of three alert ily when handled. The mother returned to the furred-out young as early as May 7, 2005. nest in less than five minutes after we left. Two lactating females were collected on 14 Since one nestbox was appropriated May and 19 June. Young, or at least small from chickadees, it was interesting that on specimens thought to be young-of-the-year, 15 August, three black-capped chickadees were taken in October and November. In (Parus atricapillis) approached the mother Oconto County a small squirrel was taken 19 partially hiding in a jack pine crevice, about 2 July, and in Juneau County as early as 1 m away, coming within 1 m, 1.5 m, and 2 m March. A nest was also observed 19 March distance all making the distraction display call, 1991. Courtship begins early, earlier than in without the usual display (see Long, 1982a). G. sabrinus, in February. There seem to be In a couple minutes they left, and the female second litters. One lived 7 years (Crandall, returned to her nest. The young were proba- 1964). On my property in central Wisconsin, bly capable of gliding away from the box.. I have seen naked young in nests in Septem- Mortality. Nocturnal predators such as ber and others haired out in October (capa- great horned owls (Errington et al., 1940), ble of “flight”) and likewise in late April, early house cats (Jackson, 1961; Hoffmeister, May, and late July. One observed litter of two 1989) and other owls and carnivores (Dolan males, naked and blind, were subsequently and Carter, 1977) prey upon G. volans. reared by the mother in a bird house (with They are occasionally injured by impaling gnawed entrance). The two were seen first 16 themselves on sharp objects while gliding. April, the lengths about 50 mm. The mother Snakes may enter tree hollows and eat them. did not move the young and usually watched Timber cutting and forest fires locally eradi- us from a nearby oak sapling as we checked cate these squirrels.

218 THE WILD MAMMALS OF WISCONSIN Table Rod-7. Some external and cranial measuremenrs of Wisconsin Glaucomys.

Counties Sex TL Tail Hind Foot Ear Gr. L Zygo br Max. t-r Glaucomys sabrinus Clark F 298 112 32 21 37.2 21.7 7.04 Forest M 265 113 32 23 36.3 20.6 6.7 Forest F 273 112 34 23 38.0 22.4 6.6 Portage M 294 118 36 21 37.2 22.1 6.9 Portage M 260 111 39 21 – – 6.8 Portage F 276 112 36 22 38.0 22.2 6.84 Oneida M 250 120 35 18 35.7 21.2 – Oneida M 240 100 40 20 35.6 – 6.74 Oneida F ?217 ?50 35 22 37.6 – 6.85

Glaucomys volans Portage 6M 212±26 92±4 28.8±2.6 16.0±3 33.8±1.7 20.4±0.7 5.86±0.2 Portage 7F 204±40 95.1±6.7 30.3±3.8 16.0±2.3 35.0±0.8 20.6±0.9 6.18±0.23

Ectoparasites include six species of fleas, about 100 feet, and usually they run around mites, and lice (Day and Benton, 1980; Dolan the trunk of a tree immediately after alighting and Carter, 1977; Scharf and Stewart, 1980). on it. This may allow the squirrel to escape a Endoparasites include protozoa, acanthoceph- following owl. There is a communal torpor of alan worms, roundworms, and tapeworms irregular periods, but no true hibernation, and (Dolan and Carter, 1977). Likely a rare inci- these squirrels are active year round. dence, rabies and a type of malaria have been Additional Natural History. Dolan and reported (Venters, 1962; Dasgupta and Chat- Carter (1977) is an excellent source for G. tergee, 1967). See account of G. sabrinus. volans. Hatt (1931) noted habits. Home Range and Density. There is little Geographic Variation. None was ob- information on populations of G. volans in served in G. volans. Wisconsin, and for nearby states the infor- Specimens examined. Total, 42. Adams mation is confusing and speculative, both for Co, Chippewa, Dane, Douglas, Dunn, Ju- movements and density. For one thing, the neau, Monroe, Oconto, Polk, Portage, Vil- young of second litters join the adults and as, Waupaca, Wood counties. earlier young in autumn to make the densities higher. In both species of flying squirrels, judging by my museum specimens, the numbers of young in September, October, and November were nearly triple the usual numbers per month. On my 1.2 acres of woods overlooking McDill Pond in Portage County, there have been occasional visits by one or two adult squirrels to feed on acorns. Numer- ous females have brought forth litters in the bird houses, but never more than two females at once, and the numbers of young were mentioned under Reproduction. Remarks. Nocturnal gliders, the small G. volans can glide rapidly for distances up to Wisconsin Beaver. By C. A. Long.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 219 Family CASTORIDAE Gray “beavers” and as a kind of animal “beaver”, e.g., “three beavers were on the bank”, or Genus Castor Linnaeus “some beaver burrow in stream banks.” Beavers Description. Beavers are large (dog-sized) rodents with brown or tawny brown color. There Holarctic, large, aquatic rodents having dor- is a dorsoventrally flattened tail, oval in dorsal soventrally flattened, nearly naked and scaly outline, usually scaly and mostly hairless, but tails, webbed hind feet and a sulcate zygo- furred proximally. It varies in size from 20 cm matic plate anterior to the zygoma. The pri- (8 inches) X 10 cm (4 inches) to 32.5 (13 inch- mary food, at least in winter, is the inner es) X 20 cm. The tail functions for fat storage bark (cambium and phloem layers) of trees, for winter, communication by slapping the wa- which are felled and gnawed, and often inter surface, heat exchange, balancing while corporated piece by piece, summer and fall, walking on land, and, of course, as a rudder in in dams and lodges. The family is ancient, swimming. The hind feet are large with webs arising in the Oligocene Epoch. The aquatic between the toes (see fig.), having the first and genus Castor is known only through the especially the second digit with a divided claw Pleistocene and Recent. (for grooming the fur). All feet are pentadactyl, the forefeet dextrous and used to handle foods, carry young beavers (“kits”), and transport build- Castor canadensis Kuhl ing materials to construct the dams and lodges. North American Beaver The underfur is gray and dense, the brownish guard hairs coarse and long. “The Beaver of Canada... surpasses the Badger in The skull is large with formidable root- size, and is the most industrious animal in less incisors (chestnut on the anterior surfac- fabricating its dwelling... Large glandular pouches, es) and rootless hypsodont cheek teeth (hav- which terminate on the prepuce, secrete a ing transverse lophs). A diastema is present pommade of very pungent odour, which is in each tooth-row. In dorsal aspect, the paired employed in Medicine... Castoreum.” — nasals, taken together, are oval, widest in the G. Cuvier, in Regne Animal, 1846. middle. There is a small sagittal crest in older specimens, and the zygomata are stout and 1820. Castor canadensis Kuhl. Beitr. z. Zool. und very deep at the postorbital process. The zy- vergleich. Anat.1: 64. Type locality from Hud- gomatic plate where the masseter muscle at- son Bay, Canada. taches is deeply sulcate. The dentary shows a 1912. Castor subauratus Taylor. Univ. California deep angular process projecting ventrally, and Publ. Zool., Vol. 10, p. 167. Type from Gray- below the lower incisor the dentary may son, Stanislaus Co., California. project ventrally somewhat as in sabre-tooth 1913. Castor caecator Bangs. Bull. Mus. Comp. cats. The basioccipital is dished, and the zy- Zool., Vol. 54, p. 513. Type from Bay of St. gomatic arches flare away from the skull. George, Newfoundland. The cheek teeth were described in Reg- ne Animal as “grinders, four in number above The name Castor comes from Greek and below, with flat crowns. They appear as kastor, meaning beaver, and canadensis re- if formed of a doubled bony fillet, exhibiting fers to the country where the type was col- one deep indentation on their internal bor- lected. Beaver is an old English word “beo- ders, and three on the outer edge above, and for” meaning brown. There is no rule about the reverse below” (G. Cuvier, 1846, 1863). the plural of beaver, interchanged more or There is a club-shaped baculum in males less. Mention of particular animals usually is tapering to a blunt or rounded distal tip, and there

220 THE WILD MAMMALS OF WISCONSIN are four pectoral mammae in females. There are The color is brown, but the tone may be 40 chromosomes (Lavrov and Orlov, 1973). chestnut, reddish brown, tawny or drab wal- Other aquatic adaptations include a nicti- nut, even black. The tail is nearly black but it tating membrane used when swimming, and the pales with age to gray. The color is nearly the closure of nostrils and ear canals under water. same overall, brighter above, and usually lack- By bradycardia and other physiological adapta- ing splotches of other colors. However, Lov- tions beaver can submerge up to 15 minutes. allo and Suzuki (1993) reported a family-re- Anal scent and castorium (one of the lated phenotype of splotched white feet in two earliest scents used in perfumes) glands are beavers. The underfur is dense, and there is a present in both sexes. These latter glands near molt beginning in summer. Prime fur devel- the groin empty with the urine to scent-mark ops for winter. piles of vegetation and mud (castors) or other The total length varies from about 1,000 sites. The anal glands are paired. The lips close to 1,200 mm (39-46.6 in), tail length 258- behind the incisors, probably an aquatic ad- 440, tail width, 90-200, hind foot length, aptation. Probably also adaptive to aquatic life 156-205, and ear length, 23-29. Cranial is the cloaca, a pouch that accommodates the measurements range up to 146 mm greatest scent glands, the urogenital openings and rec- length, 108 zygomatic breadth. The weights tum (this cloaca is not to be confused with range usually to about 30 to 40 pounds but any reptilian cloaca). It is difficult to identify occasionally in Wisconsin exceed 100 pounds the sexes because of the cloaca. Other char- (= 45 kg). Schorger (1953) reported large acters are listed by Jenkins and Busher (1979) Wisconsin beavers weighing 39.4, 36.5, and and Hill (in Chapman and Feldhamer, 1982). 35.6 kg. Some measurements are listed in Table Rod-8. Recently Ucker (1994) submitted a photo of a carefully weighed beaver from the Peshtigo River, near Crivitz, weighing 120 pounds (54.5 kg), a new weight record. Walter Klukas was the trapper. The total length is nearly as long as the height of the man shown with the beaver. Dental Formula. I 1/1, C 0/0, P 1/1, M 3/3 = 20. Geographic Distribution. Before 1800, beavers occurred throughout what is now Wisconsin and the Upper Peninsula of Mich- igan. Native Americans used them for food Skull of Castor canadensis. (Baker, 1983). The Caucasion Americans almost eradicated them in this region. Since 1901, the wildlife conservation agencies in Wisconsin and surrounding states protected surviving beavers. The species made a remarkable recovery, not only in Upper Michigan and Wisconsin, but throughout most of the United States. The beaver is uncommon in the prairie counties of western Wisconsin, and does not seem to be a permanent resident on most is- Webbed hind foot of beaver. lands in Lake Superior or Lake Michigan. It is

TAXONOMIC ACCOUNTS / ORDER RODENTIA 221 abundant on Outer Island, in the Apostles. Beavers may visit islands from time to time, and may take up residence on some of the larger ones. From the 1970’s until the present the beaver has been regaining lost territory, except in urban areas, and may occur in all the western, southwestern, central, and northern counties. It lives near people, if left alone, so long as food and water are available. Status. Beavers build lodges and dams, modifying local environments in Wisconsin marshes, swamps, and stream valleys. Of-

Maps of geographic distribution for Castor c. canadensis and C. c. michiganensis (north of west-east line above Green Bay) in Wisconsin and North America.

222 THE WILD MAMMALS OF WISCONSIN ten this distinctive habitat is called a “bea- (Greenman, 1966). Legal protection began ver dam community.” It often affects the in 1901, and was practiced more stringent- distribution of many organisms, including ly by 1905 (Jackson, 1961). About 1919, waterfowl, trout and many wetland species a study on the status of beaver made by (Knudsen, 1962). Certain fishes, amphibi- Hartley Jackson and George Wagner deter- ans, birds, mammals, and some inverte- mined that beaver were an asset to Wiscon- brates and plants lived in the beaver dam sin, despite some flooding of some roads communities, which changed from im- and agricultural land and cutting of desir- poundment to meadow as beavers felled able trees. Transplanting began in the early aspens, willows, and other deciduous trees 1920’s. Some beaver from Wisconsin and before moving on to new habitat, leaving Upper Michigan were sent to Pennsylvania the dams to decay and drain. Beaver also and New York. In 1936, beaver occurred lodge in lakes and dig bank burrows in river in 50 counties, and by 1954 in 56 coun- banks. During the 1700’s-1800’s in Euro- ties. Trapping was permitted since 1936 in pean and American markets, beaver fur be- some counties. Today the declining value came stylish, especially beaver hats, and the in fur results in higher numbers of living demand for North American beaver pelts beavers, and the activities of people cause led to the early exploration and exploita- many of them to consider beavers as nui- tion of the American West, as well as many sances. Over 3,000 trappers annually trap of our states, provinces, and territories east- an estimated 60,545 beaver, mostly in Coni- ward. By 1900, along with land-use by the bear traps (Dhuey, 1995). invading Caucasians, the demand for fur and Benefits to humankind (Hill, 1987) in- overtrapping led to the extirpation of bea- clude water storage (much more important vers in many vast regions of America. Bea- in the arid West, but of some Wisconsin use vers were thinned out in southern Wiscon- in flood control), erosion control of local sin, but beavers were present in Wisconsin gully formation, the fur (depending on the and upper Michigan. styles and demand), and even as a food (for The beaver inhabited the Northwoods some people). Beaver are prey for wolves for centuries. Not only are beaver bones and some other predators, and the beaver most abundant of mammalian remains at dam community benefits many other ani- most archaeological sites, a burial with cop- mals and plants. Aesthetic appreciation for per implements contained a beaver robe nature observers is important. Formerly castorium was used in perfume making and as a medicine. Jackson (1961) discussed the formation by beavers of humus soils and many low prairies (or at least meadows) in Wisconsin. Over long periods of time, silt accumulates in impound- ments of small valleys in the highland, established successively by beavers working their way toward the valley floor. These beaver meadows in sum have helped produce much rich farm land in valleys of wooded regions across the Lodge of beaver. Where dikes and waterways are entire northern half of North America (Ruede- damaged, culverts dammed up and roads flooded, thousands mann and Schonmaker, 1938). of nuisance beaver are transported to wilder places (if they are unoccupied by other beaver, and budget and time allow). On the negative side, beavers cut down C. Long, The World Book. ornamental trees, nibble on shrubs, burrow

TAXONOMIC ACCOUNTS / ORDER RODENTIA 223 in dikes, and cave off stream banks. They room with its floor several inches above the occasionally follow a row of corn in a field water level. The beavers do not work in eating the shoots. Their extensive flooding groups, each tends to work alone (Novak, of low cultivated areas by dams that are 1987). At least two entrances are below sometimes huge and complex may cause con- water. Beavers also dig burrows into river- siderable damage. banks, occasionally combining one kind of Long famous for its fur and its impor- dwelling with the other. To swim freely from tance in the exploration of America, scientif- one pond to another, beavers occasionally ic and popular literature are replete with arti- dig canals (not over 6 m in my experience), cles on the beaver. Only the white-tailed deer some in Upper Michigan were mentioned may rival the beaver in literary volume. In by Agassiz (1913). Jackson (1961) gave the Canada, the beaver is a famous symbol found length of some as 300 feet. (I suggest that in some endearing movies, books and arti- water flow helped ditch such a long trench.) cles, and on coins and seals. In northern lakes where the water is sur- Habitats. Being aquatic, the beaver in- rounded by granitic and steep banks, where habits various bodies of water: lakes, riv- no burrow could be dug, and the water is ers, ponds, streams, dammed up brooks and too deep for a lodge sited out in the water, ditches, marshes, swamps, and impound- the lodge may be constructed on a steep ments. Dams constructed are often large rock ledge at the water’s edge, and not be and complex, and impound water over wide separated from the shore. If there are no areas (a few ponds exceed 10 acres). They stream banks, a lodge must be constructed. are deep enough to provide security to the Water may be impounded by one or more beaver colony. Thus, wild areas may be al- dams. These may be small or taller than a tered by the activities of beavers, usually man’s height. changing the land to a wetland. Food trees Alexander Agassiz, the famous natural- are necessary in beaver habitat. When used ist son of the great naturalist Louis Agassiz, up the beavers depart, leading to another studied beaver for two years in Upper Mich- ecological change, from the beaver dam igan. He was also busy developing a copper community to that of a sedge or grassy mine. He wrote to another scholar on bea- meadow. Not only do many animals and vers, Lewis H. Morgan, about his “repeated plants associate with the beaver in its man- opportunities” in Upper Michigan, to study ufactured environments, but several kinds beavers collecting chips, observing habits, even have used their lodges for their homes, dams, and even canals [italics mine]. He es- including water shrews, wolves, and bob- timated some colonies were over 900 years cats (see accounts of those species). old, that some ponds exceeded 40 acres in From August to September the beaver area, that the associated beaver meadows colony works on lodge building and dam- were up to two mile-sections in area, and ming of water. The sound of flowing water that beavers were active all winter, often wan- reportedly is a stimulus to build dams. The dering far from the lodge, with many tracks behavior is seen in juveniles, but they may going “in all directions”. not do construction the first year. Downed In Ashland County a dam was construct- trees, branches, mud, stones, and other ed 12 feet high and 40 rods long (Barber available material, even corn stalks (Robert 1919). Here a giant lodge was built 16 feet Henderson, personal correspondence) are high. The lodge may be as low as one meter used. The beaver digs, pushes and shovels but is usually up to two meters high. As many (lifts) mud and other materials into place, as eight to ten beavers may occupy a large building a dome-shaped roof over a large lodge. As it is built, over a single room, the

224 THE WILD MAMMALS OF WISCONSIN surrounding mud is excavated, so that the between 3 and 4, occasionally with more water near the lodge deepens as the lodge kits, sometimes 6, reportedly 10. (With only grows higher. The temperature inside on cold four mammae probably about 6 is the max- winter nights is warmer than outside, and imum number of young). Longley and Moyle the lodge temperature is not so extreme as (1963) found litter size in Wisconsin as 5.3 the air temperature outside. The kits are born embryos. The more food available the high- inside the lodge or bank burrow. er is the litter size. Population density af- Foods. The primary and essential foods fects fecundity (Payne, 1984, Boyce 1974). are inner bark of aspens (Johnson, 1983), Large mothers tend to have large litters. The paper birch or willow. Aspen is much pre- young beavers are born completely furred ferred. Other species of trees (at least tree out, in various shades of color much as in parts) are eaten. Various water plants, buds, the adults, with eyes partly open and teeth and other terrestrial plants are eaten on oc- partially erupted. They require care but casion, such as water lilies (personal obser- seem rather precocial compared to most vation) and even poison ivy, sedges, tubers rodents. They leave the lodge with the moth- and roots, corn, and acorns. Conifers regu- er in 14 days, and males may bring in leaves larly are eaten, especially young white pines for them to eat in 14 days. The infants range in winter and spring (personal observation, in size from 340-630 g, with members of over 20 years). Novak (1987) mentions that small litters tending to weigh more. Some conifers are not essential, but beavers may southern populations tend to have small lit- girdle small pines and eat the pitch. Coproph- ters, but there seems to be no evidence prov- agy has been observed. ing a relation to temperature or latitude Small trees are easily cut down and (Payne, 1975; Osborn, 1949; Hill, 1987). hauled to the water, where the beaver eats in Beaver often live up to 13 years, occasion- relative security. Trees about 3-5 inches in ally to 24 (Nowak, 1991). diameter are often hung up by other trees, The young are weaned at about 8 weeks and much of their food is unused (Johnson, of age, some, perhaps, by three months, but 1983). Trees larger than that usually fall to they feed readily on plant material after about the ground and may be used. Trees leaning one month, and some, reportedly, as early as over the water will fall into it when cut, and 4 days. The young are suckled about nine are readily used for food or construction. times a day. The young fondle the mammae Beavers may wander 100 feet or so from as they drink during 5-10 minute periods. water looking for food. Food may be trans- Young-of- the- year (i.e., kits) apparently ported through canals. In autumn and win- do not breed their first winter, but some year- ter, the Wisconsin beavers cache branches lings may be active the second winter. Breed- under water for food in the season when the ing is listed as 21 months for both sexes (No- pond or stream is locked in ice. vak, 1987). Young are usually not driven out Reproduction. Beavers tend to be mo- of the lodge in autumn, but many depart be- nogamous, and to maintain offspring of sev- fore then to establish lodges of their own. eral ages in the lodge. Usually a single fe- Mortality. Humans cause much mortali- male in the lodge conceives. She breeds gen- ty from trapping, 13-80 percent (Novak, erally in January or February, into March. 1987), and transplant some beaver to new Beaver often copulate in water. Castors (i.e., habitats (Boyce, 1974; Payne, 1975). Floods scent mounds) are made at this time for ter- and hunger kill some beavers naturally. About ritorial reasons. The gestation lasts about 107 the only natural predators are wolves, which days, with birth occurring in May or June, prey often on beavers in summer, especially and sometimes later. The mean litter size is when deer are scarce (Hill, 1987). Jackson

TAXONOMIC ACCOUNTS / ORDER RODENTIA 225 (1961) lists the great-horned owl and goshawk 100 females, or to a greater proportion of as predators. Occasional predators are river females (Gunson, 1970; Payne 1975; Wood- otters, even mink, coyotes, red foxes, bears, ward 1977). Apparently there is a prepon- bobcats, and lynx. Black bears preyed on bea- derance of males in subadults but more fe- vers in the Apostle Islands (Smith et al. 1994). males in adults. (See account of the black bear). Tularemia is Briefly, this curious and historically im- a disease of beaver, which also affects hu- portant mammal, with such a complex family mans; it is apparently water borne and also life and clan territory, may be said to be crep- transmitted by ectoparasites and direct con- uscular and nocturnal. It is often seen in Wis- tact. It is caused by the bacterium Francisella consin in the daytime. The young live with tularensis (= Bacterium tularense). Tulare- the adults, and sometimes other young are mia sometimes causes important epidemics also there, during the winter months. Move- in beavers. A variety of protozoans and worms ments (Hill, 1987) include colony activities, are internal parasites (Jackson, 1961). Exter- wandering by some adults not really under- nal parasites are scarce but include ticks (Ix- stood but ascribed to those beaver that lost odes, and peculiar beetles (Platypsyllus and mates, and regular dispersal of 2-3 year olds Leptinillus). Occasionally, beavers are killed from a colony to form new colonies elsewhere. by falling trees they have cut themselves. Jack- A colony is a group of beaver consisting of son (1961) mentions one such casualty, and I the adult pair, yearlings, and kits. They use have a photograph of another. Beavers often and defend a common food source and a com- fight, especially males, and seem to attack the mon water supply, and often a common sys- rump and tail. Although wounds are common, tem of dams. The territory is marked at least mortality may not be much affected. in some areas by scent mounds called “cas- Home Range and Density. Movements tors”, small piles of mud, vegetation, and se- have been as extensive as 238 km, but aver- cretions of castorium from the castorium aged only 7.4 km in 472 translocated beavers glands and probably scent of the anal glands. that were recaptured (Knudsen and Hale, This scent is known to cause disturbance in a 1965). Dispersal reduces density of beaver colony when emitted by foreign beavers. colonies, but the main controlling factor is trap- Warning sounds are made by slapping the tail ping (Boyce, 1974). As lowered food quality on the water surface, but vocalizations are few, may deplete beaver densities, depleted food consisting of low whines, whistles, or whim- caches may cause starvation and wandering in pers. The young learn to slap their tails in the winter and spring, leading to greater mortality water at about 2-3 weeks of age. The beaver by predation. Density is usually expressed as tries to run from enemies like any other ter- colonies per given area, say 10 km2 square, or restrial mammal, but it can walk erect using a given length of stream (i.e., 1 per 10 km of the tail as a counterbalance, carrying mud and stream). Colonies are located in the fall by the other objects in the forepaws. It swims both presence of food caches. Data published out- at the surface and below it, usually kicking side Wisconsin vary from about 0.3 colonies alternately with a hind foot. It is active all year, per 10 km2, sometimes three times that in good but often confines its activities in winter to habitat, and along streams about the same. the lodge, and to the food cache. On warm Remarks. A relative of Castor canaden- nights it may be out on the banks. They sel- sis that formerly lived in Wisconsin was the dom attack and bite, but they often hiss and giant beaver Casteroides, mentioned in the threaten. account of prehistoric mammals earlier. The behavior observed within a lodge is The sex ratio varies from place to place, highly interesting. Prior to birth of young the from about even numbers, from 95 males to entire family was observed to clean the lodge

226 THE WILD MAMMALS OF WISCONSIN and spread leaves and herbs over the floor. at least on the character of coloration, it be- In this endeavor the family works as a group. came necessary as a taxonomist to draw some The family observed the birth, and the large kind of boundary between this race and bea- male and another beaver formed a triangle vers observed in central Wisconsin. Including with the mother to enclose and warm the kits. some beavers mounted long ago, I have seen The mother ate the placentas. Juveniles and only tawny brown beavers in central Wiscon- adults lick the rather precocial neonates, and sin. They occasionally attain great size (ex- carry them away from water. Juveniles clean ceeding 100 pounds). I cannot follow Jack- the lodge, bringing in new vegetation. The son (1961) in assigning all Wisconsin beavers mortality of kits is surprisingly low in most to the Michigan race. That judgement gives studies, even of young-of-the-year. Few bea- credence to Hoffmeister’s (1989) provisional ver live to ages as old as 10 years, but occa- classification, and removes an Hoffmeister- sionally to 20 (see Novak 1987). Jackson example from along the Illinois state Additional Natural History. Jenkins and boundary (Table Plan-1). Busher (1979) reviewed biology for the bea- Most specimens examined, unfortunate- ver. A classic study was made by Warren ly, are skulls (the pelts were sold), and the few (1927) on the beaver in Yellowstone Park. skins do not allow any definition of subspe- Geographic variation. Two geographic cies boundary. Not many skins are available races occur in Wisconsin. They differ signif- in the Smithsonian collection either, and that icantly in color, but not much by size or pro- is probably why Jackson (1961) ascribed the portion. From my observations, and by use entire state of Wisconsin to the dark race. That of the literature, I conclude that the dark Upper Peninsula beavers are dark brown, Michigan beaver is of medium size, whereas nearly black, is well known in the fur busi- the beavers in most of Wisconsin are medi- ness. It is a common saying there that Iron um to large. Probably southern beavers av- County beavers are the same as Upper Pen- erage larger, but possibly have lived longer. insula beavers. John Olson, a DNR fur spe- That being the case, one can assume the cialist at Park Falls, believes some there are physiological and reproductive traits of the nearly black, some “light”, and most brown. two kinds hardly differ. There is little evidence Mr. Russ Kilker, the Gruenwald Fur Compa- that either kind has been altered racially by ny, believes most beavers in Wisconsin are transplantation and introductions in the last brown, but become lighter or “reddish” near- 70 years. Certainly some of the beavers in ing the Illinois boundary. Wisconsin are large, even huge, but the size Kelly Thiel, a beaver trapper for U. S. differences are really not distinctive in the Department of Agriculture (damage control), variable family groups. traps beavers that are nuisances and those Dr. Al Gardner, recalling his field work often found along trout streams. He has rein Seney National Wildlife Sanctuary in Up- portedly handled 1,200 beavers yearly since per Michigan, told Robert Fisher and me, 1988, in northern Wisconsin. He has a ra- “they [beavers] are not only dark, but to me tional perception of beaver variation based they looked black.” Across the Upper Penin- on much experience. Although he has seen sula, and ranging into northern Wisconsin no beavers from Iron County, he observed (where all the beavers examined by Jackson, beavers black, almost black, and occasional- 1961, had been trapped) the beavers are dark ly dark brown in Bayfield, Douglas, and Vil- in color, a soft almost black tone varying from as counties. Some are blackish in Ashland otter brown to a charcoal black (not glossy County. Between Rhinelander and Merrill he black, as in skunks or bears). Convinced of believes a line could be drawn, south of which the validity of the northern geographic race, in Lincoln and Marathon counties beavers

TAXONOMIC ACCOUNTS / ORDER RODENTIA 227 are usually brown, and lighter (golden brown), Description. A dark colored, medium- occasionally an odd color of reddish-brown. sized beaver closely resembling the nominate Based on his impression, which agrees with race. See Geographic Variation above. that of Olson, and is consistent with my few Specimens examined. Total, Wisconsin observations, I draw a provisional boundary 5. Bayfield, Vilas counties. across the state, noting that individual varia- Other records. Bayfield Co.:Taylor’s tion and family variation broaden the per- Creek (Jackson 1961). Douglas Co.: Loval- ceived zone of intergradation between the lo and Suzuki (1993). Upper Michigan (USNM races. The line separates a distinctive north- specimens). Luce, Alger, Chippewa counties. ern dark race from southern and western beaver populations. Table Rod-8. External measurements and some weights Marked variation evident in four pelts in for beavers in Upper Michigan and northern Wisconsin. the Madison collection show these traits: Por- tage Co. Dark reddish brown; Dane Co. pale Total tail hind foot lengths golden tan; Grant Co. in molt (March) with US 170561 Taquahmenaw River 58 lbs worn golden brown pelage and dark reddish US 247288 Chicogan Lake 1015 300 175 — brown new fur; and Green Co. very dark, al- US 243780 Michigamme 1055 350 175 32, 40 lbs US 243788 Michigamme 1030 290 175 35, 30 lbs most blackish, but head golden brown. US 247287 Salmon Trout Cr. 960 300 170 —, 30 lbs US 236596 Grandview, WI 1050 350 180 35, — US 236595 Grandview, WI 965 365 170 30, 35 lbs Castor canadensis canadensis Kuhl US 247318 Gresham Lake 1025 390 170 —, 40 lbs

See above synonymy of the nominate race given for the species. Type Hudson Bay. Table Rod-9. Fur harvest data of aquatic furbearers in Description. The critical trait for identifi- Wisconsin. After Dhuey, 1994, 1995, 2000, and personal cation is color, nearly always bright chestnut- communication. golden brown, and seldom a dark walnut brown. See geographic variation above. Species Year Harvest Average $ Pelt Value Remark. In some other states, introduced Price beavers of subspecies differing from the one Muskrat 1999-2000 286,212 — — extirpated there has led to some hesitation in Muskrat 1996-1997 160,747 4.15 $667,100 using geographic races at all for beavers Muskrat 1995-1996 108,046 2.35 $253,908 Muskrat 1994-1995 325,396 2.21 $719,125 Specimens examined. Total, 31. Buffa- Muskrat 1993-1994 304,741 1.86 $566,818 lo, Dane, Grant, Green, Lincoln, Marathon, Mink 1999-2000 16,921 — — Portage, Price, Taylor, Waupaca, Wood Mink 1996-1997 21,333 20.50 $437,326 counties. Mink 1995-1996 8,545 12.08 $103,223 Mink 1994-1995 32,500 11.84 $384,800 Mink 1993-1994 22,086 16.15 $356,689 Beaver 1999-2000 — — — Castor canadensis michiganensis V. Bailey Beaver 1996-1997 30,184 20.44 $616,960 Beaver 1995-1996 25,012 16.46 $411,697 1913. Castor canadensis michiganesis V. Bailey. Beaver 1994-1995 86,574* 14.45 $1,250,994 Beaver 1993-1994 61,339 15.23 $934,193 Proc. biol. Soc. Washington, 26: 192. Type Otter 1999-2000 2,794 — — from Tahquamenaw River, 5 mi. above falls, Otter 1996-1997 1,233 44.66 $62,462 Luce Co., Upper Michigan. Otter 1995-1996 1,233 44.47 $54,837 Otter 1994-1995 4,615 48.76 $225,027 Geographic Distribution. Northern Wis- Otter 1993-1994 3,412 57.30 $195,507 consin and Upper Michigan. See Map. *6,201 beaver taken in supplementary harvest.

228 THE WILD MAMMALS OF WISCONSIN Family GEOMYIDAE Gill Genus Geomys Rafinesque Pocket Gophers Eastern Pocket Gophers

The fossorial pocket gophers are sciuromor- The North American rodents Geomys are ph rodents that have, with their closest rela- represented in Wisconsin by one species. tives, the Heteromyidae, evolved paired fur- lined cheek pouches with openings outside the mouth. This adaptation for hoarding food Geomys bursarius (Shaw) in the burrow was different from internal Plains Pocket Gopher cheek pouches of deer mice, chipmunks, ground squirrels, hamsters and other rodents Geomys bursarius bursarius (Shaw) in having the external openings. By some macroevolutionary and probably embryolog- 1800. Mus bursarius Shaw. Trans. Linnaean Soc. ical change (Long 1977) the new kind of London, 5:227. Type from Upper Mississippi cheek pouch was immediately adaptive, be- Valley. cause the seeds transported and cached in 1817. Diplostoma fusca and D. alba Rafinesque. the burrow were dry whereas the internal Amer. Monthly Mag., 2:44. Types from Mis- cheek pouch brings all the seeds into con- souri River Region. tact with saliva in the mouth. This loss of 1825. Ascomys canadensis Lichtenstein. Abb. k. saliva to the seeds is a water loss, and in Akad. Wiss. Berlin, for 1822: 20. Type from deserts small mammals cannot afford to lose “Canada”. water. The change in foraging from stuffing 1829. Geomys bursarius: Richardson. Fauna Bor.- the seeds in the mouth instead into the near- Amer., 1:203. by fur-lined pouch (in fact, into the extend- 1957. Geomys bursarius wisconsinensis Jackson. ed and inverted corner of the mouth) per- Proc. Biol. Soc. Washington, 70: 33, type from mitted the “wrong side out” pockets to per- Lone Rock, Wisconsin. sist and improve in form. No alternative microevolutionary scenario makes any sense. The name Geomys means mouse of the earth. The specific part of the binomen bursarius means pocketed. Thus, the two characteristics, fossorial behavior and external cheek pouches are described by this Latinized name. The common name is plains pocket gopher, or merely pocket gopher or gopher. The word “gopher” is used in the vernacular way, for in much of the Midwest it is a term used for the 13-lined ground squirrel Sper-

Figure showing cheek pouches, large forefeet and grooved incisors of Plains Pocket Gopher. Figure of Geomys bursarius. C. Hart Merriam.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 229 mophilus tridecemlineatus. In central Wis- The dorsal color is drab reddish brown to consin, where no pocket gophers are to be light chestnut brown. There may be white on found, the people use the word gopher for the tip of the tail. There are at least two molts ground squirrels, without knowledge of the annually, and two may be taking place simul- common name 13-lined ground squirrel or taneously. The pelage is short, smooth and has the former name “13-lined spermophile.” a brassy sheen in sunshine. There is white-spot- Description. In Geomys, unlike other ting on nose and throat on numerous speci- pocket gophers, there are two grooves on the mens (including seven from this collection) from anterior surface of each upper incisor. The the Brule region (Jackson, 1961). median groove is deeper. A shallower but dis- The penis bone is a simple rod with tip tinct outer groove is parallel to the median expanded slightly laterally. The length is 10- one. In some of my specimens there are three 12 mm (Burt, 1960). There are eight mam- grooves, although the third is faint. Hoffmeis- mae. In mothers the pubic symphysis is enter (1989) found the same pattern in some larged (Hisaw, 1925). The chromosomes specimens in Illinois. There is a diastema on number 2N = 72, with a fundamental num- each side of the dentition, but there are up- ber of 72 with 68 acrocentrics (Hart 1978). per premolars present, each comprised of two The males are 10-15 percent larger than joined columns. The molars are almost loph- females, and reportedly at least 20-25 per- odont with transverse enamel loopings. The cent heavier. See tables Rod-10-11. rostrum is long, with the nasals extending pos- Dental Formula. 1/1, C 0/0. P 1/1, M teriorly to the level of the zygomata. The zy- 3/3 = 20. gomata are wider in their transverse breadth Geographic Range. The range of the anteriorly than behind, and the tympanic and pocket gopher is confined to the western third lambdoid expansions broaden the posterior of Wisconsin and unknown on the Upper part of the skull (the occiput is nearly straight Peninsula. If a straight line were drawn from across). Thus, the skull is rather angular, and Ashland in Bayfield County to Lone Rock and the profile in lateral view is of a quite flat- Gotham in southern Richland County, only tened braincase. Most of these specializations the record from southern Price County would are fossorial adaptations, general ones of all lie east of that line. There are no records south digging mammals, or unique ones seen only of the Wisconsin River in southwestern Wis- in pocket gophers (such as the slender tail consin, for the River apparently has halted which is a tactile organ as the gopher backs the southward dispersion. up in its burrow). Far to the southward extending across central Illinois into Indiana, is a large, black pocket gopher G. b. illinoensis. Apparently the Mississippi River prevented the species from invading Illinois along much of its length, but in southwest Illinois the crossing was made and the bridgehead expanded across the state with marked differentiations occurring in col- or and larger size. The pocket gophers of western Wiscon- sin likewise seem to have crossed the Missis- sippi (and St. Croix) rivers somehow. They occupied the sandy loam valleys extending their range eastward along the river courses, Skull of Geomys bursarius. but were barred from Grant and Iowa coun-

230 THE WILD MAMMALS OF WISCONSIN ties, and other counties in the southwest region, by the wide Wisconsin River. At Goth- am the habitat changes abruptly to jack pine savanna and across the river southward are deciduous and sycamore bottomlands which have not been invaded by Geomys, nor by red squirrels for that matter (see account of Tamiasciurus). The Wisconsin pocket gophers are tan-brown and, although some are as large as muskrats, the Wisconsin races are smaller in size than the Illinois race. The dis- similarity suggests two separate invasions, different evolutionary histories, and long separation of the two eastern stocks.

Maps showing geographic distribution of Geomys bursarius in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 231 Status. The Wisconsin pocket gopher is saliva on thousands or hundreds of hoarded not common but in no peril of eradication. seeds and stem parts (Long, 1977). Something of a nuisance, its mounds and bur- The pocket gopher lives most of its life, rows tear up agricultural fields, and there is actively in winter or summer, either night or some eating of truck crops and shocked grain. day, in elaborated tunnel systems. The bur- It tends to dwell in friable soils on well-drained row is not especially deep, except in winter ridges or embankments, and in grassy or weedy when the frost enters the ground. The tun- open areas. There the soils are sandy, with nels may descend below three feet depth. Even little clay or “gumbo” soil present. Pocket go- then, the pocket gopher tunnels to the sur- phers prefer soils where tubers and succulent face, burrows under the snow, and leaves leafy plants such as clover and dandelions are evident above ground, in warmer seasons, the available for food. Living in extensive burrow cores of plug materials for meters in length. systems, it is no surprise they avoid flood plains They were deposited inside tunnels but were and marshy areas where the burrows are sub- eventually exposed by the thawing ice and ject to flooding. Heavy farming of the land, or snow. The tunnel system usually follows a the presence of a badger can eradicate a pop- main linear tunnel, with numerous extensions ulation of pocket gophers. Rather peripheral to either side that are usually plugged with to the farmer’s sphere of activity, they are not light soils. The behavior to plug a side tunnel often harmed. Gopher traps are available in opening leads to their capture by gopher hardware stores for local problems. traps. These are set into the main tunnels Jackson (1961) states that pocket go- beneath a large and usually sandy mound, with phers inhabit “waste ground,” and therefore sand thrown more toward one side. Opening cannot be called a menace (i.e., a pest). In the tunnel by use of a shovel soon results in a such situations it often does more good than response by the gopher to plug the opening harm by turning over the soil and mixing veg- and push the trap out. The gopher sets off etable matter with it. the trap when it contacts the trigger. In some Habitat. The plains pocket gopher dwells side tunnels, which may be plugged with soil, in friable well-drained soils, usually sandy the food cache is stored. The gopher may loam, in open meadows, pastures, roadsides, extend its head outside a tunnel to the sur- railroad rights-of-way, old fields, cultivated face to tear off leaves and stems, which are fields especially clover and alfalfa, cutover or cut into suitable lengths to transport into the burned over land, and occasionally in lawns food tunnels. Roots and tubers are eaten be- or gardens. In Wisconsin, the gophers are low ground. The main burrow over a long usually in fairly close proximity to streams, time period may have required removal of because that is where the sandy soils that the some seventy cubic feet of earth. Obviously gophers use as immigration routes into the burrow systems vary with terrain, time of ex- hilly countryside. In sandy areas, the proxim- cavation, soil composition, and the individual ity of water also permits the establishment of whims of the fossorial homebuilder (Smith forbs and legumes, dandelions, clover, alfal- 1948, Downhower and Hall 1966). McLaugh- fa, and other succulent plants on which go- lin (1951) reported a system 125 feet long phers feed. Gophers are not found in the wet with at least 57 mounds. No studies have been or clay soils, or in dense root masses in wet- made on tunnel systems in Wisconsin. Ap- lands or along streams, and according to Jack- parently the usual times that tunnels are son (1961) they cannot swim. Their external shared are when young are with the mother cheek pouches are not only adaptive in hoard- or two gophers are mating. ing foods into their tunnels, but are an arid- The nest is reportedly a mass of cut environment adaptation restricting the loss of stems, observed at a depth of 2 feet, measur-

232 THE WILD MAMMALS OF WISCONSIN ing about 7 X 3 1/2 X 6 1/2 inches. The cut the Mississippi and St. Croix rivers in Wis- stems are about one and a half inches in consin, contained no embryos. On 7 May a length, comprised of common grasses in the female (UW-SP 7699) was lactating and con- area (Smith 1948). McLaughlin (1951) also tained 5 embryos 8-12 m in crown rump described a nest and a food cache. length. Another (UW-SP 7700) contained 2 Foods. Hardly anything is known of food embryos 10 mm long. A third female con- habits of Geomys in Wisconsin. They doubt- tained no embryos. There are usually three less eat alfalfa, clover, and dandelions. Other or four neonates (young) in a litter, but the foods of similar nature are probably eaten, number varies from 2-6. The newborn young and grasses and tree roots may be eaten. are blind and naked, about two inches in Bones were gnawed in the nest of a gopher length and weighing six or seven grams. The studied by Smith (1948). skin is pinkish. There are two furrows or Reproduction. According to Jackson grooves (Anlagen) for the future external (1961), who cites no studies, mating takes pockets at the corner of the mouth. Each fur- place in the spring, sometimes by late March, row is about 5 mm in length. The young may but usually during April or May. The male may emit squeaks after birth. They are nursed at tunnel into a female’s burrow or travel over least 10 days, perhaps until two-thirds grown, the surface of the ground in search of a fe- at which time they may poke their heads from male. A specimen from nearby Allamakee Co. holes to forage. Eventually they tunnel away Iowa was pregnant (3 embryos) on 8 April to make new burrows on their own. There is 1969. The embryos were 11 mm in crown- a single litter each year, and young-of-the-year rump length. Females taken in April, east of cannot breed. McLauglin (1951) found male and female cohabiting a burrow system on February 9, in Illinois, of a different subspecies of Geomys bursarius. Bailey (1929) reported embryos between April 12 and May 22 in Minnesota. Hoffmeister (1989) gives the gestation as about one month, but a Kansan gopher in captivity gave birth 51 days after capture (Sud- man et al. 1986). In Wisconsin, judging from my few records, it lasts about a month (late April to late May). Mortality. Little is known about the enemies of Geomys in Wisconsin. A young or above-ground gopher may be caught by any carnivore, hawk, or owl. Owls frequently regurgitate remains of Geomys in owl pellets. Jackson (1961) suggests that weasels catch them, and in nearby Minnesota the pocket gopher is the primary prey of the badger (Lampe 1976). It is odd Jackson made no mention of the badger as a predator, probably due to a paucity of evidence. Bull snakes prey on them (Hisaw and Gloyd 1926). Farm- Burrow system of Geomys bursarius. Courtesy E. ers often rid their fields of pocket gophers. Raymond Hall.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 233 Home Range and Density. The home the Mississippi and St. Croix rivers, the pock- range is approximately the same size as the et gophers in Wisconsin have not differentiat- burrow system, because the pocket gopher ed much. They do not approach the distinct- seldom leaves the system for ordinary wan- ness of the isolated population in central Illi- derings. Extensions of burrows result in dis- nois, which is predominantly black. Pocket persal movements. Jackson (1961) thought gophers are highly variable from place to place, those might be “many miles” but this distance leading to the naming of many populations as seems doubtful. geographic races. In Wisconsin specimens I There may be male-female occupancy in have examined there is marked variation with a burrow in February or March. A solitary age and males are larger than females. mother will occupy a burrow with her young The northernmost gophers do not cross until they are weaned. Usually males outnum- the Brule River northward, but range along ber females and likely wander more. In the its southeastern shore. They have extended UW-SP collection the percent of females is their range across the divide of the Mississip- 60 percent (N=25). There is little informa- pi and Superior drainage in Bayfield County, tion available about home range and density but not by much. According to Jackson (1957) in Geomys in Wisconsin. In mild winters these northern gophers differ from southern mounds are pushed up near Osseo until populations in several cranial characters: The Christmas. Fresh mounds are evident in late premaxillary bones reportedly are wider and March and early May. I have seen one core less concave posteriorly, almost flattened or of dirt left by melting snow, about 3 m in even slightly convex. The rostrum is report- length at 3 mi. W Mondovi, April 20, 1996, edly wider also, and the edge of the premax- and another April 24, 1997, at the same illary bone is nearly straight anterior to the place. One core was 2 m in length. zygomatic arch. A narrower distance sepa- Remarks. The excavation-function was rates the posterior ends of the nasals and the described long ago by Merriam (1895) as pick- ing and excavating with the incisors while the forefeet both dig and shove loose dirt under Table Rod-10Rod-10. External measurements of Geomys bur- the body. The hind feet also move the dirt sarius. All specimens judged adult, but some are old adults. posteriorly, and if the gopher does not suc- Males supposedly exceed females in size, but some observed ceed in moving beyond the accumulation it females were exceptionally large. Measurements from Jack- turns in the burrow and with both wrists joined son (1961) are marked with an asterisk. The northern specimens precede the southern. together under the chin, pushes the dirt back with the palms of the hands while moving with Locality Sex&N Total L. Tail Hind Foot Wts g the hind feet. Thus, the dirt is discharged from NW M2 292 85 35.5 310 the nearest opening forming a “little hillock”. Wisconsin* (260-325) (75-95) (34-37) (270-350) This hillock is called a gopher mound. Go- F2 230, 279 68, 85 31, 34 — pher mounds usually reveal the presence of Douglas Co. M4 260 73.3 33.8 — the secretive gophers. The mounds are not &Drummond (249-269) (66-80) (33-35) — F2 259 70 33 — usually circular, as are mole hills, because the (249-270) (70-70) (32-34) — gopher throws the dirt to one side. They are 3 W Strum M4 313 87.5 37.8 — more likely comprised of sand. It is odd that Bell Center (277-375) (80-92) (34-37) — this species is so asocial, when some other Cochrane F1 242 67 36 — burrowing rodents find such benefit in com- Osseo F1 224 69 31 183 1/2 N Chippewa F1 283 90 35 — munal living. Falls Geographic Variation. Even though iso- 2 S Galesville F1 255 66 31 — lated in the western counties of Wisconsin by 1 E Gotham F2 260,260 63,73 31,31 260.6 g

234 THE WILD MAMMALS OF WISCONSIN Table Rod-11Rod-11. Some cranial measurements of Geomys Subfamily Murinae Gray bursarius, males from Gordon, Strum, Bell Center, and Lone Rock, and females from Chippewa Falls and Lone Rock. “What[ever] moss is to the reindeer, what[ever] Greatest L Zygomatic br Cranial br Max. t-r grass is to the buffalo, the Mouse millions of the North are to all the northern carnivores, from Bear 52.3 31.2 26.5 8.9 52.7 33.5 28.8 8.8 to Blarina. When we shall have fully worked out the 53.0 33.3 29.4 9.7 life history of each of these species, I believe we 52.6, 31.2, — — shall learn that the whole of that vast, beautiful, 54.1* 31.7 — — important, and specialized production that we call 54.4 33.8 29.6 8.9 “carnivora,” rests on a broad simple basis of 42.5-44.5* 25.4-26.2 — — Muridae, that in turn rests on the grass, that rests *Occasionally the nasals extend farther than the incisors, and on the earth. We shall for each of these flesh-eaters occasionally the supraoccipital extends farther posteriorly than the write, “It sometimes eats this and sometimes eats condyles. that, but by far the greatest bulk of its food is Mice.” — Ernest Thompson Seton, Food of the Marten, in adjacent frontal bones are likewise narrower. “Furbearers” This distance is only about one-half as broad as it is long, instead of noticeably wider. How- Myomorph rodents (i.e., those with a ever, the characters did not work in my spec- small infraorbital canal traversed by a slip of imens (Table Rod-11), and if they did the dif- the masseter muscle) are those lacking preference would be trivial. Jackson mentioned molars, and as defined here consist of mice the color is not “appreciably” different, but and rats (species of Peromyscus, Reithrod- my southern specimens are slightly paler. No ontomys, Microtus, and allies all formerly significant geographic variation was observed. called cricetids), as well as the introduced Specimens examined, Total 42. Bay- house mouse Mus musculus and Norway rat field, Buffalo, Burnett, Chippewa, Crawford, Rattus norvegicus, which have sparsely Douglas, Dunn, Eau Claire, Jackson, Mon- haired, scaly tails. and hard palates extend- roe, Richland, Trempealeau counties. ing posteriorly behind the last molars.

Family MURIDAE Gray Rats and Mice, Some are Quite Nice

These are myomorph rodents (see remarks under Rodentia) and include herein the common North American mice known previously as Cricetidae (the genus Cricetus is not native to North America). Some workers use Sigmo- dontidae, which excludes Mus and Rattus. Our Peromyscus and Microtus are the commonest mammals in Wisconsin, and are wide-spread.

Skulls of Peromyscus leucopus, P. maniculatus (dorsal and ventral views, note pinched rostrum), Reithrodontomys megalotis (dorsal and ventral, note grooved incisors), and the vole Pitymys ochrogaster (3 views a, b, c, note zig-zag molars). After E. R. Hall and D. Hoffmeister.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 235 Genus Reithrodontomys Giglioli Reithrodontomys megalotis (Baird) Harvest Mice Western Harvest Mouse

Often found in sheaves, haystacks, and round 1858. Reithrodon megalotis Baird. Mammals, in bales of hay associated with harvest, the mice Reports Expl. Surv... 8(l): 451. Type locality give credence to their name by bending grass between Janos, Chihuahua and San Luis stems over to cut off the heads of seeds. These Springs, New Mexico tiny mice are delicate in form, brownish above, 1893. Reithrodontomys megalotis: J.A. Allen. Bull. clean white below, bright-eyed, and their up- Amer. Mus. Nat. Hist., 5:79. per incisors are each grooved on the anterior face for the length of the crown and much of the root (see figure above left). The skull is Reithrodontomys megalotis dychei rounded and smooth, and has slender zygo- J. A. Allen mata. The incisive foramina are large and hardly separated by a thin septum. The outer 1895. Reithrodontomys dychei J.A. Allen. Bull. wall of the infraorbital canal is a broad, deli- Amer. Mus. Nat. Hist., 7:120. Type from cate sheet of bone. The tail is thinly haired, Lawrence, Douglas Co., Kansas. distinctly bicolored, and usually elongate in 1914. Reithrodontomys megalotis dychei: A.H. relation to the total length. The feet are ex- Howell. N. Amer. Fauna, 36:30. ceptionally small, smaller even than those of 1944. Reithrodontomys megalotis pectoralis Han- Peromyscus maniculatus bairdii (see Mea- son. Field Mus. Publs., 564, Zool. Ser., 29- surements). Of 18 species recognized in North 205. Type from Westpoint, Columbia Co., America, there is but one in Wisconsin, and Wisconsin. Hoffmeister and Warnock, Trans. it is confined to the southern half of the state. Illinois Acad. Sci., 47-162, 1955, and other taxonomists have regarded the Wisconsin mice as inseparable from R. m. dychei. The difference, a pectoral spot, is inconstant in Wiscon- sin harvest mice.

Reithrodontomys translates as “grooved- tooth mouse”, which is meaningful. The specific part of the binomen megalotis means big ear, which is less meaningful (although the ear is prominent). The subspecific part of the trinomen, dychei, honors L. R. Dyche, the founder of the University of Kansas Museum of Natural Histo- ry, and the collector of the holotype. Description. See account of the genus above. This small, clean-appearing mouse, with grooved incisors and tiny feet, can be identified by several cranial characters and small body. The skull, described above in characters for the genus, has total length less than 22 mm. The baculum is a minute rod. The penis and baculum resemble those larger structures Harvest mouse. By F.B.S. in Anthony. in Peromyscus. There are six mammae (one

236 THE WILD MAMMALS OF WISCONSIN pair pectoral, two pairs inguinal). Unlike 48 fa, sandy prairies, and grassy edges of marshes chromosomes seen in all Peromyscus, the (hayfields) so far only in the southern half of harvest mouse has 2N=42 (Shellhammer the state. See Map. 1967; Webster and Jones 1982). Extra chro- Status. The harvest mouse is rare in Wis- mosomes (up to four tiny chromosomes) are consin. Nowhere is it common. The prairies reported in some specimens. Grooved inci- on which it lived in southern and southwest- sors resemble those in the pocket mouse Per- ern Wisconsin were themselves greatly dis- ognathus (found in Minnesota, but unknown turbed, although some mice have established in Wisconsin) and jumping mice (Zapodidae). themselves in what remains and in old fields, For comparison with long-tailed Zapus, Mus pastures, and savannas. This mouse may have musculus, which resembles a dirty-colored invaded the state from the west, from Minne- harvest mouse, and Peromyscus manicula- sota, or the south or southwest, Illinois or tus bairdii, see accounts of those species. Iowa, about the turn of the 19th century. The Some external and available cranial mea- mouse had extended its range deeply into surements of Wisconsin specimens are as fol- central Wisconsin (Long 1970), but no spec- lows: Portage Co.: Female 127-61-16-15; imens have been taken recently (in spite of greatest length 19.9, zygomatic br. 10.3, int. some diligent efforts) for over 20 years. The br. 3.25, maxillary tooth row 3.7, length nasals threat of land use destroying prairies and even 7.5, wt. —; F 141.5-63-19-14; —; M 139-68- old field habitats is great in southern Wiscon- 12-13-10.0 g. Waupaca Co.: 127-50-18-14; sin (e.g., Dane County). This beautiful and —; Trempealeau Co.: 120.5 (112-130), 57(50- harmless mouse should be monitored by state 60), 15.8 (15-17), 12(12-12), N = 4; 19.6, authorities, and transplants of harvest mice 10.1, 3.2, 3.4, 7.1 (N = 1). Monroe Co.: Fort into some of the newly established prairies McCoy 129-60-16-11, 9.0g.; 21.1, 10.6, 3.3, would be interesting. 3.2, 7.0. A pregnant female from Sauk Prairie Birkenholz (1967) suggests the harvest measured 120-51-14-X and weighed 10.3 g. mouse occupies seral, open habitats. That Dental Formula= I 1/1, C 0/0, P 0/0, might explain why some populations decline M 3/3=16. over time. Land use might also explain the Geographic Distribution. The harvest declines, as might unknown causes such as mouse occurs in grazed fields, stubble, alfal- long-term climate change. Conservationists will study these problems. Habitat. The harvest mouse is found in grassy areas with or without brush or trees present, seral or disturbed fields, grazed pastures, marsh lands, hayfields, and sandy ridges. Grasses may include brome, bluegrass, bluestem, foxtail, Panicum, as well as various forbs such as goldenrod and smartweed (Birkenholz 1967; Hanson, 1945). Birkenholz found that they preferred habitats with grasses about 18 inches in height, whereas Hoffmeister (in an overgrown orchard with blackberries) found them in vegetation as high as five feet. Webster and Jones (1982) mention grassy and weedy habitats such as pastures, meadows, fence-rows, fallow fields, and Family and nest of harvest mice. Lloyd Sanford. borders of fields. In southern Wisconsin, Sv-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 237 endsen (1970) caught this mouse in fair abundance in 1967-1968, in old field habitats, especially in autumn, in Phlelum, Agropy- ron, and Panicum grasses, the legumes Les- pedeza and Trifolium, the forbes Aster, As- clepias, Aplopappas and Solidago (comprising 72 percent of the total plant species). The grasses Andropogon, Elumus and Setarus, the blackberry Eubatus, and the composites Tragopogon and Taraxacum made up 23 per cent of the plants. A few woody plants and grasses made the remainder.

Maps showing geographic distribution of Reithrodontomys megalotis in Wisconsin and North America.

238 THE WILD MAMMALS OF WISCONSIN The nests are balls of grass stems and Remains are often identified in owl pellets. leaves set in a field above ground. They are Parasites include Trypanosoma, a cestode, occasionally placed in an abandoned bird’s acanthocephalan, nematodes, fleas, chiggers, nest. They may be sited two feet above the mites and lice (Webster and Jones, 1982). ground. The nests are often lined with milk- Some parasites are mentioned by Jackson weed or thistle down (Schwartz and Schwartz (1961). Mumford and Whitaker (1985) list 1981). Small openings are found at the base parasites from Indiana. of the nest. Harvest mice also live in burrows Home Range and Density. Schwartz and (Birkenholz 1967), especially in winter. Schwartz (1981) and Kaye (1961) report the Foods. No food studies have been made home range of two females as one half to a on Wisconsin harvest mice. Elsewhere the diet little larger than one and one half acres. In is primarily seeds, many of which are cached Illinois over a three year span, Birkenholz for winter. Much herbaceous vegetation is also (1967) found the density to fluctuate normal- consumed. Insects (as much as 22% Lepi- ly from 4-12 per acre. In central and western dopteran larvae) are eaten (Whitaker and Wisconsin, seldom can even two be taken at Mumford 1972). the same locality. Svendsen (1970) found the Hall (1959) quotes Henry Fitch stating population density much higher in southern that this mouse consumed seeds of foxtail, Wisconsin, in 1967-1968, reported as 18 switchgrass, Indian grass, false redtop grass, mice per acre (45 / ha) during the fall sea- brome, and side-oats grama. Harvest mice sons. He reports this amounts to 0.046 ani- climb weeds, legumes and grasses day and mals per trap night. In spring only 0.012 an- night to harvest seeds. imals per trap night were taken. There were Reproduction. A single pregnant female four times as many mice in autumn. Hansen collected in the Sauk Prairie contained 4 (1945) found a maximum density of 2.4 per embryos. The mice breed only during the acre in foxtail-smartweed cover. growing season. Some data are summarized Remarks. Information on this little mouse in Webster and Jones (1982) and Jackson is so scarce, I failed to find a petite vignette (1961). The gestation period is 23-24 days, from some famous naturalist to charm the possibly longer. Newborn are naked, blind, reader, to make it known this is no common and helpless. They weigh only about one gram mouse. The old masters had so little informa- each. The eyes open in less than two weeks. tion they could seldom identify any species of Fur grows out in about one week. The young harvest mouse. Some failed to note the are weaned in about 24 days. The litter size grooved upper incisors, and one said they is about four (2-6 or 7). In Indiana, Mumford never ate insects. Cory (1912) had no speci- and Whitaker (1982) found 30 pregnant fe- mens from either Wisconsin or Illinois. Em- males to average 3.8 embryos (2-6). There met T. Hooper was an authority on murid are rarely as many as 8 or 9 embryos (Long, mice, including Central American harvest 1962). Adult size is reached in five weeks. mice, but said nothing for my purpose. Hall The young mice may mate by six weeks. A (1955) mentioned that they could travel many second litter may be produced 24 days after yards through the high stems of grasses with- the first, according to E. R. Hall. The life span out coming to the ground, and described the may be 18 months. feet as efficient for seizing and grasping grass Mortality. Nothing is known about pre- stems because the toes of the forefeet flex dation on harvest mice in Wisconsin. I saw against the palms making a grasping hand. I what I think was a harvest mouse “treed” on offer my own vignette: The harvest mice seem a tall weed by a large snake. Prairie carni- to be mice of the grasslands, indeed living in vores and raptors take them when possible. unity and harmony with the community of

TAXONOMIC ACCOUNTS / ORDER RODENTIA 239 grasses themselves. Regarding this ecologi- dan Swamp 2. T21N, R8E, Sect. 4, 1. Sauk cally integrated relationship of grass and har- Co.: Sauk Prairie 1 UW Wild. Ecol. Trem- vest mouse, the poet Walt Whitman might pealeau Co.: 2 mi. N Gaylesville 4. Waupaca have agreed, perhaps, in his Leaves of Grass, Co.: 1 mi. N Iola 1. singing the grassland was “as the father to bosom held his son.” Harvest mice travel through the tops of the tall grasses, make nests Genus Peromyscus Gloger of the fine grass stems, line the nests with the softest fibers available from milkweeds and The American deer mice, formerly called thistles, eat grass seeds, prey on grasshop- “white-footed mice” (a name now used for pers, and make or use runways extending Peromyscus leucopus) inhabit most of North through grass. One pet I had for a while made America, often with two or more co-existing its nest from a ball of cotton fibers, and like (i.e., sympatric) species. They are usually abun- my captive Peromyscus spent much time dant and are among the most abundant of grooming itself and playing about in the cage. mammals in the ecosystems of America. Such The pelage is a pale brownish gray, the belly abundance is true in Wisconsin, where three and feet purest white (dusky at the hair bases). interacting Peromyscus are found. Owing to The dainty form seemed especially charming, their manifold life-activities and high popula- but there was no petting this pet. When open- tion densities, even their great biomass as ing the door to clean the cage one had to be forage for predators, they are indeed biolog- careful. The sprightly mite leaped suddenly ically important. William Henry Burt (1948), onto an arm, out the door, up the curtains, famous mammalogist, mentioned that the and then a “merry chase” ensued. forest deer mouse was exceptionally beauti- Hall and Kelson (1959) mentioned that ful. That mouse occurs in northern Wiscon- of the harvest mice the plains species is the sin, and the other two Wisconsin deer mice most widespread. Likewise, this Wisconsin are also beautiful. The three mice resemble subspecies has extended its range deep into one another and are not easily identified with- this state. The nominate race R. m. megalo- out reference to several cryptic but diagnos- tis occupied the vast range of basins and high tic characters (see below). prairies west of the Rockies, and the Great This genus in the past was known as Plains race R. m. dychei occupied the vast Hesperomys Waterhouse or Sitomys Fitz- prairies east of the mountains from Alberta inger. The type species for Peromyscus is in Canada to eastern Arkansas. Crossing the Peromyscus arboreus, a synonym of Peromy- Mississippi River only with one salient, the scus leucopus noveboracensis. The deer race may have invaded northwest Illinois and mice resemble murine rodents such as Mus did find southwest Wisconsin. Eventually it by having long tails (as long as a third or even found its way into central Wisconsin. more than half of the total length). There are Additional Natural History. Webster and minute scaly annulations of the tail clothed Jones (1982) studied the plains harvest mouse. over with fur. The ears are mostly naked and Geographic Variation. One race dispersed prominent. Internal cheek pouches are more northward or northeastward into Wisconsin. or less developed. The relatively large eyes, Specimens examined. Total 37. Colum- thin-walled cranium lacking conspicuous ridg- bia Co.: Portage 1. Dane Co.: York Twsp. es, small infraorbital canal, roughly triangular 1. Various localities 20 UW. Monroe Co.: Fort and bounded by a bony plate of the anterior McCoy, Tl7N, R3W, Sec. 20, 1. Fort McCoy part of the zygoma, all characterize these 2 UW. Portage Co.: 5 1/2 mi. N Stevens mice. The dentary is elongate with short coro- Point 1. 5-5 1/2 mi. E Stevens Point 2. Jor- noid process, molars are low crowned and

240 THE WILD MAMMALS OF WISCONSIN tuberculate (when unworn), and the upper The two long-tailed Peromyscus resem- incisors are smooth. ble one another, and often are in similar habitats. The juveniles are not identifiable in such situations, although they soon attain adult Key to the species of Peromyscus characters (e.g., large ears in P. maniculatus). See the Key above for general traits. 1 Hind foot usually 17-18 mm, to 19 mm, Long and Long (1993) used bivariate and dis- tail short, usually less than 64 mm in criminant analysis of adult mice in northern length, upper parts brown darkening and central Wisconsin, and on islands in Lake dorsally nearly to black ...... Superior and Lake Michigan. They found geo- ...... Peromyscus maniculatus bairdii graphic variation in P. maniculatus suggest- 1’ Hindfoot usually 20-22 mm in length, ing that the name gracilis in Wisconsin seems tail longer than 64 mm, dorsal fur brown inappropriate (see account beyond of P. without much mid-dorsal dark tone, but maniculatus). In their analyses the best char- wuth reddish chestnut brown or dusky acters were ear length, length of rostrum, and fawn (yellowish tan-brown) ...... 2 length of tail. Other characters varied from 2 Ears 17 mm or longer, tail about 80-83 place to place, but incisive foramina length, mm in length and pencillate (tufted), whis- cranial depth and breadth (previously given high kers long (the upper vibrissae may be ex- weight by taxonomists) were least reliable. tended behind the ear pinna, the lower vibrissae are nearly as long and white), dorsal fur yellowish in tone or brown ..... Peromyscus leucopus (Rafinesque) ... Peromyscus maniculatus maniculatus 2’ Ears less than 16 mm in length, tail less 1818. Musculops leucopus Rafinesque. Amer. than 77 mm, never pencillate, and sel- Monthly Mag. 3: 446. Type from Kentucky dom well-haired, whiskers “normal” Pine Barrens. length (some upper vibrissae may be ex- 1895. Peromyscus leucopus: Thomas. Ann. Mag. tended even behind the ear pinna, but Nat. Hist., series 6, 15: 192. lower vibrissae obscure), dorsal fur often slightly reddish or brown ...... Although Hoffmeister (1989) uses the ...... Peromyscus leucopus name of the nominate race P. l. leucopus for Illinois white-footed mice, I follow Osgood Blair (l940) provided a classic of popula- (1909) and numerous other workers in rec- tion dynamics for Peromyscus in southern ognition of the race P. 1. noveboracensis Michigan. King (1968) reviewed biology for (Fischer). Hoffmeister’s rationale was that this genus. Wolff (1989) described social be- some Illinois variants were similar in size to havior in Peromyscus in Kirkland and Layne’s Kentucky mice. Cory (1912) ascribed south- (1989) review of biology. Wolff (1985) dis- ern Illinois mice to the nominate race, but cussed ecology in Peromyscus. most Illinois mice to the race noveboracen- The prairie deer mouse P. m. bairdii is sis. Cory mentioned larger size, longer fur, small, with a short, thick tail. The fur is dark and paler upper parts in noveboracensis. The mid-dorsally; the short tail (< 65mm) and hind venters are purer white. He also referred the foot (usually 17-18 mm) easily identifies this Wisconsin white-footed mice to this race. mouse in the field. Young mice, in gray pel- Osgood (1909) had analyzed over 300 mice age, have a less attenuate tail than seen in from the nominate race, restricted by him to the other long-tailed species. In the laborato- the “Lower Austral” life zone (p. 118), and ry the short skulls identify adult P. m. bairdii. over 2,000 of noveboracensis. Hoffmeister’s

TAXONOMIC ACCOUNTS / ORDER RODENTIA 241 observation that mice in northern Illinois are mings, nor as long as in long-tailed P. manic- not readily distinguishable from the southern ulatus) and a dusky brown or chestnut brown mice may be understood by Osgood’s com- dorsum with pure whitish underparts (often ment that ideally the holotype of leucopus tinged buff or cream). The tail may be “dis- would have been selected from Louisiana and tinctly bicolored” (dark brown above and whit- for noveboracensis from the middle of New ish below) or in many specimens sparsely and England. Intergradation confuses comparisons thinly haired, so as to appear “indistinctly bi- from Kentucky, Illinois [and Wisconsin]. Os- colored”. The contrasting colors more or less good describes well the geographic variation merge when they meet. This latter character in white-footed mice from this Wisconsin- was not original with Burt (e.g., Audubon and Michigan region, and the northern Illinois Bachman, 1842), but Burt made it famous in mice are noveboracensis. his field guide (Burt and Grossenheider, 1952) and it does not always work. When the tail is indistinctly bicolored, which is often the case, Peromyscus leucopus that color pattern identifies P. leucopus. noveboracensis (Fischer) From P. m. maniculatus, which P. leu- White-footed Mouse copus resembles closely, the white-footed mouse has a shorter tail (65-78 mm), shorter 1829. [Mus sylvaticus] noveboracensis Fischer. Syn- whiskers (vibrissae), the lower ones are never opsis mammalium. P. 318. Type from New York. white, and shorter rostrum of the cranium with 1830. Cricetus myoides Gapper. Zool. J. 5: 204. an anteriorly pinched pair of incisive forami- Type from a place between NewYork and Lake na. The ear pinnae are shorter (smaller) than Simcoe, Ontario. in the race maniculatus, and are similar but 1842. Mus michiganensis Audubon and Bachman. larger than in the short-tailed P. maniculatus J. Acad. Nat. Sci. Philadelphia. P. 304. Type bairdii. The tail lacks a pencillate tuft of hairs from Erie Co.: Ohio. This name was applied and as a rule is more sparsely haired than in P. to Peromyscus maniculatus bairdii, and prob- m. maniculatus. In lateral profile the skull is ably to Mus musculus. See their synonymies. short and shows an arched cranium. 1901. Peromyscus leucopus minnesotae Mearns. Adult P. leucopus differ from the small Proc. Biol. Soc. Washington, 14: 154. Type P. m. bairdii in having larger feet (19-21 mm from Fort Snelling, Hennepin Co., Minnesota. length, which are also wider or thicker. In comparison to the house mouse Mus muscu- The name Peromyscus means “little lus, P. leucopus lacks a buffy or brownish mouse” but the meaning of the prefix Pero is unknown. The name leucopus is Greek for “white foot.” The name noveboracensis is Latinized, intended to mean “from New York,” based by Fischer on Pennant’s New York “variety.” Eboracum refers to the English area now called York, Eboracum to the Romans, later changed to the name of a Viking warrior Jorvik, which eventually became Yorwik, and finally York. The word novum means new and “ensis” means from this place. Description. A long-tailed deer mouse having medium length whiskers and ear pin- Photo of mother white-footed mouse suckling young. nae (not short as in the voles and bog lem- 1900. By Dugmoor.

242 THE WILD MAMMALS OF WISCONSIN belly (a whitish belly is seen in some Mus), and the eyes are less protruberant and not set forward. P. leucopus differs from the harvest mouse Reithrodontomys megalotis in having smooth (lacking grooves) upper incisors, and it is decidedly larger in body dimensions. The baculum is a simple rod slightly curved (Hooper, 1958). The number of mammae is six, with at least four pectoral teats functional. The chromosome number is 48 (Singh et al., 1966). In more detail than mentioned above, the color of juveniles is gray above, whitish be-

Maps showing geographic distribution of Peromyscus leucopus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 243 insula of Michigan, anywhere in forest edge, mature deciduous forest, and brushy habitats. This species is not confined to such habitats, has been steadily dispersing northward from southern Wisconsin, but is excluded from many of the dense northern coniferous and mixed hardwood-conifer forests. It is excluded from some habitats in prairies, swamps, and marshes. Centuries ago this mouse invaded deeply into Wisconsin (Jackson 1961). Jackson (1961) reported the geographic range as approximately the southern three- fourths of the state, with the marginal records then mapped at the tip of the Door Peninsu- la, and from southern Oconto, Price, and Washburn counties. He believed that the species had not followed the land use by human- Overlap and movements of long-tailed Peromyscus Fine kind as aggressively as some species had (e.g., line is margin for P. m. maniculatus; it receded and has relict populations in the south. Heavy dashed line shows extensions Spermophilus tridecemlineatus or P. man- of range for P. leucopus. The dotted line shows range for iculatus bairdii). Burt (1948) had listed only leucopus mapped by Jackson (1961). Long (1996). one county record from the Upper Peninsula (in Menominee Co.). As can be seen from the low, as in other juvenal Peromyscus. The map, the white-footed mouse has greatly ex- adults of all Peromyscus are indistinguishable panded its range since 1961, even invading in color of the ear pinnae (being slate gray) Washington Island (Long and Long 1993; and eyes (shiny black). The dorsum varies Long, 1996), and several far northern coun- depending on the fresh or worn condition of ties, and also has increased its range on the the pelage; worn pelage is dark brown (the Upper Peninsula of Michigan (see Baker, plumbeous or lead gray basal parts of the hairs 1983). Agricultural fields and roadsides are show through and the bright tips are worn important dispersal routes (Cummings and away) whereas fresh pelage (as soon as the Vessey, 1994). bright tips grow out) is nearly russet or chest- Peromyscus leucopus is aggressive, nut. Rarely is such reddish pelage seen in P. adapting to human land use and dispersing maniculatus, never in P. m. bairdii (which is northward, as is evident from the mapping olivaceous walnut brown or even grayish of past (Jackson, 1961; Burt, 1948) and brown, and darker along the mid-dorsum). present records. Northward dispersion also The feet are whitish. The adults molt annual- was evident on the Lower Peninsula of Mich- ly, in summer. igan (Hooper, 1942) and in New York and See Key for measurements and compar- Connecticut (Miller,1893). There seems to isons with other long-tailed deer mice. Ac- be competition between long-tailed P. man- companying figures show weights in males iculatus and P. leucopus, or perhaps the and females. land use is becoming more favorable to leu- Dental Formula. DF = 1/1, C 0/0, P 0/ copus. That P. leucopus ecologically re- 0, M 3/3 = 16. placed the long-tailed maniculatus along a Geographic Distribution. See Map. This zone of contact in Wisconsin and Michigan, species may be expected in suitable habitats roughly along the so-called Tension Zone” throughout the state and on the Upper Pen- ecotone, is based on zoogeography (Long,

244 THE WILD MAMMALS OF WISCONSIN 1996). Particularly useful are relicts of “gra- eats grain and garden products, but also is cilis” (= P. m. maniculatus) in the south, beneficial in eating insects and weed seeds. and the pattern in Door County, where P. The white-footed mouse doubtless com- leucopus occupies the Peninsula and long- petes with other granivores and nut-eaters. By tailed P. maniculatus occurs on numerous burying acorns it aids forestation, but may re- offshore islands. Either replacement occurred place P. m. maniculatus when they occur sym- on the Peninsula, or P. maniculatus invad- patrically. It may invade the rural habitations ed all the islands and missed the Peninsula. of people and cause a little damage. One po- In Clark County, at the northwestern margin of known range, the two species maintain separate but similar habitats in the west and are sympatric in the east. On Washington Island, P. leucopus invaded in 1987 (Long and Long, 1993) and soon increased in numbers (Long, 1996), while P. maniculatus decreased dramatically in the forest habitats (see Fig.). Since then the two species fluctuated irregularly (Long, 1996). Other studies show P. maniculatus is better adapted to severe cold (Wolff, 1986; Wolff and Hurlbutt, 1980; Pierce and Vogt, 1993). Since the observed competition ensued on Washington Island, in the past six relatively mild winters, neither species seems dominant. No significant differences were seen in diet, parasites, or effects of predation. Status. This mouse may be the most abundant wild mammal in Wisconsin. It occurs in most counties. In the forests and savannas it seldom does any damage to humankind, and indeed is a valuable member of predator food chains. In agricultural settings it occasionally

Scatter plots for long-tailed Peromyscus groups Above Graph showing frequencies before and after P. leucopus bivariate for ear and tail lengths. Below Canonical variates invaded the home of P. maniculatus, Washington Island. of discriminant functions. Long and Long, 1993. 2’s = central After Long (1996). Wisconsin P. leucopus.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 245 tential problem is the possible transmission of on the northern and northwestern lakeshore disease either directly or by its ectoparasite bluffs. Stah (1978;1980), Wolff and Durrand hosts (ticks, fleas). These include plague, Lyme (1986) and Wolff and Hurlbutt (1982) report disease, Rocky Mountain spotted fever, and that the two co-exist in the same woodlots, rarely the recently discovered Hantavirus. and that P. maniculatus dwells more often Habitat. See general comments above in in the trees. In such a place, P. leucopus likely the section on Geographic Distribution. The would eventually win out, with its better base white-footed mouse is a forest-edge species, for ground foraging. It is more likely to use but is also found in many diverse forest and ground dens. In cold, snowy and boreal for- woodland habitats (beech-maple, oak-maple, ests (spruce, fir, tamarack, etc.) the long tailed oak-hazelnut, jack pine, oak-sycamore). Mice Peromyscus maniculatus seem dominant thrive especially in the deciduous woodlands (Pierce and Vogt, 1993; Wolff and Durr 1986; and brushy areas (Getz, 1961; Long and Long, 1996; and others). See account of P. Long, 1993; Long, 1978). In fields, the mouse maniculatus (Table Rod-12). prefers the brushy thickets, invades the grass- Occasionally found in burrows or ground lands as brush and saplings ecologically sup- crevices, this scansorial mouse usually nests plant other prairie species, and thrives in on or above ground and sometimes in trees brambles, vines and thickets so long as they (Horner, 1954; Stah, 1980). The nests are are not too wet. The species ranges into ri- constructed of fine plant fibers, downy or fi- parian woodlands where Clethrionomys and brous material, feathers and cloth, and even its stump-deadfall habitat are found, onto rabbit fur on occasion, sited in a mass of sticks, floodplains (where the white-footed mice may weed stems, leaves and grasses. The nest is thrive, or seem absent even when the species made in abandoned bird nests, hollow logs, is abundant nearby), on sand barrens and sa- or branches of trees, crevices in rocks and vannas, in hedgerows, barn lots and pastures, stumps, and studs of man-made walls (Nichol- weedy prairies, disturbed farmlands, even in son, 1941; Jackson 1961). Eisenberg (1968) cultivated fields. Also it is found at wooded described the stereotyped behavior of nest rocky outcrops and ravines where creeks may building. The nest measures as much as 10 or may not be present. These refugia protect inches (= 254 mm) in diameter, with a depth them from deep frost (Long, 1973). of about 7 inches, and having an entrance Getz (1961) is the only worker I know of hole on the side. Male and female may jointly who found them ranging into grass-sedge build the nest. Feces and especially a gooey marshes, although I have taken them at the strong urine in winter may foul the nest and edge of marshes where different habitats are necessitate making another one. nearby. In winter the open, wind-swept fields, Foods. The chief foods of white-footed where the snow is thin, is death for white-footed mice are seeds of forbs and grasses, nuts, pine mice, which survive in tree cavities, bird hous- seeds, and insects. In spring and often at other es or similar protected places in their winter times berries seem to color the finely minced nests. They may seek shelter in barns and hous- stomach contents. White-footed mice hoard es, basements, and garages. I twice observed caches of seeds or nuts both below and above hoards of acorns on the engines of cars. ground in summer and autumn (Heithaus, On Washington Island, where now this 1981). Whitaker (1963) and Grahame (1929) species is found with P. m. maniculatus, the listed insects as foods, larvae of moths, beetles white-footed mouse tends to occupy higher, and even butterflies, as well as sawflies, wasps, dryer woods and to congregate in the vicinity and hornets. Sometimes these mice eat car- of oaks (which are not common trees there). penter ants, both the adults and the eggs (Long, The P. maniculatus climb better and persist 1996). Grass and weed seeds, clover, and fruits

246 THE WILD MAMMALS OF WISCONSIN such as grapes or cherries are eaten (not hoard- The bimodal pattern may be more ap- ed), as well as some fungi (Gosling, 1977; Getz, parent in southern Wisconsin, and is seen in 1961; Whitaker, 1967). southern Michigan (Burt, 1940) and central Reproduction. In central Wisconsin the Illinois (Long, 1968). Another example of a carry-over mice from the winter produce two bimodal curve is given by Svendson (1964) successive litters usually and most of the adults for northern Kansas. In the southern latitudes perish. In late summer and autumn the young there are more exceptions to the rule that of the year produce one or two litters. The carry-overs perish, and indeed breeding oc- litter size (N=31) in central Wisconsin is ap- curs in every month of the year. But the basic proximately 4.77±0.18 (range 3-7) so that a pattern is bimodal, with a cessation of breed- female is capable of reproducing nearly 20 ing in mid-summer. offspring if she lives a full year, approximate- When winter’s deep frost is deep in the ly 9.5 in each year. Observed frequencies of ground, as it is in central Wisconsin when numbers of embryos was as follows: 3 em- snow cover is thin (occasionally the frost drops bryos, 4 females; 4, 7; 5, 13 (mode); 6, 0; to 2 m depth), then most breeding is by young- and 7, 1. Counts of placental scars were com- of-the-year in the next late summer and fall. parable. The testes of the male enlarge in With so many mice dying before spring and spring and produce viable sperm. Long (1973) early summer, the few young that are pro- showed that even the carry-over males tend duced may thrive in the open habitats. This is to perish after breeding in the spring. an example of seasonality and reproductive Long (1973a) explained the so-called adaptation. When snow cover (17-18 inches, “unimodal” peak of summer reproduction deeper in snowdrifts and in the North) pre- seen in northern populations of white-footed vents deep penetration of killing frost, the mice combined two subsets— the carry-overs breeding peaks in spring, but hardly any breed- and young-of-the-year mice. These are basi- ing adults survive through summer. Juvenile cally the same population subsets breeding in frequencies are likewise bimodal, with highs the bimodal pattern farther south (Stickel and appearing in late summer and again in early Warbach, 1960, Long, 1968, and others). fall. Long (1973) also found that the central The two periods of breeding are compressed Wisconsin mice fall into a cline of decreasing together in the northern latitudes because the litter size southward, providing an example season is shorter (the cold season is longer). of Lord’s Ecological Rule (1960), describing From one subset through the second subset, larger litters of northern, non-hibernating both testis function and lactation continue small mammals. In northern latitudes P. leu- through the breeding season, but the number copus shows increased mean litter size, al- of births declines in mid-summer (see fig.). though the number of litters may be fewer. No good reason has been found to explain this rule, because selection for large litters apparently necessary in northern latitudes, indeed, would as likely increase survival also in the south (where no reason can be found for the litters to be smaller). Large litters may be unfit to some extent, if they cause any mortality or problems for the reproductive females. Thus, theoretically any adverse effect of mortality of mothers, may balance with Figure showing subsets of unimodal frequency distribution for body weights in Peromyscus leucopus. Y = the adverse effect of small litter size, so that young of the year, n above bars. Long, 1968. the latter is confounded in the north and the

TAXONOMIC ACCOUNTS / ORDER RODENTIA 247 former in the south (Spencer and Steinhoff, combination of diurnal nursing time and sea- 1968). (But why would hypothesized mortal- sonality, favoring larger litters in the north ity in southern females having large litters not with small litters favored southward. be adverse also in the north?) Mating begins in February with gestation In birds the time spent foraging is longer about 22-25 days (Svihla, 1932). In central in northern populations because the photo- Wisconsin it usually begins a little later, in period is longer in summer there (Lack, 1968). March, and ends in September. Lactation may The food gathered will support a larger brood continue into October. The female shows a size, whereas in southern latitudes the time post-partum estrus and those that live can for foraging limits brood size. There might be breed again. Nursing the previous litter seem- a parallel cause for this gradient, and photo- ingly slows gestation of the new litter, extend- period may well be the cause. But nocturnal ing gestation to about 37 days (Hill, 1972). parent mice do not forage for food in the long- At birth the newborn mouse weighs about 1.5- er day in the north. Long (1973) hypothe- 2.0 g, and is naked and blind. The fur devel- sized the time spent nursing and caring for ops rapidly, the ears come erect, and after the litters in the nests during long daytime about 12-14 days the eyes open. The young periods might support larger litters in the are weaned in about 3-4 weeks (Layne, 1968). north, whereas shorter photoperiods limit Young-of-the-year first enter the populations them in the south. Obviously some correlate in Wisconsin in June. with latitude makes sense because the litter Mortality. A predominant cause of mor- size changes so regularly southward in the tality in central and northern Wisconsin is the same race (Long, 1973). One bothersome aforementioned lack of snow cover and con- question, does the time spent nursing during sequent deep frost (Long, 1973). Neverthe- the day also last longer at higher elevations less, in central Wisconsin, where soil frost depth (where again we find litter size is larger)? Con- is greater than anywhere in the state, P. leu- siderations of the elevation factor suggest sea- copus is the dominant small mammal. The sonality, for mouse populations. If a season- shrubby and savanna vegetation of the central ality hypothesis (Lindstet and Boyce, 1985; sands plain possibly allows P. leucopus to and others) is correct explanation, at least in thrive. The populations fluctuate with food part, then reduced density dependent com- abundance too, including mast failures, but the petition, following a winter of starvation and fecundity quickly replaces the losses. Preda- high mortality, might lead either to the evolu- tors that eat white-footed mice are many, in- tion of increased body mass or in this case cluding mostly nocturnal species because the fecundity. P. leucopus is seldom out in the day. Owls Deep, killing frost, as well as hunger, in- that take them include barn, saw whet, screech, creases mortality at the northern limits of long-eared, barred, and great-horned. Appar- range in P. leucopus. By creating open hab- ently the great gray owl does not take them itats in spring, frost and mortality favor in- normally (see accounts of Synaptomys and creased fecundity (since an increase of body Microtus). Carnivores that prey on white-foot- mass was not observed). Could seasonal low ed mice include coyotes, foxes, weasels, density-open habitat conditions in the North- skunks, raccoons, house cats, badgers, and land favor, let us say, in about 10,000 years mink. Sometimes the adults of P. leucopus and probably less (i.e., since the Wisconsinan and P. maniculatus eat young Peromyscus, glaciation), an increased litter size, and could presumably not their own. I have seen an exit also favor those mice that produce two ample of infanticide in June on Washington spring litters in quick succession? This author Island, Wisconsin, where the two species are believes Lord’s Rule is explained best by a found together (Long, 1996). Shrews and

248 THE WILD MAMMALS OF WISCONSIN snakes eat white-footed mice, and they appear lines again. At peak abundance in autumn, as in the diet occasionally of kestrels, harriers, many as 65 percent of traps contain mice. About and other hawks (Fitch and Bare, 1978; Ering- 60 percent trapped are males. In some habitats ton, Hamerstrom and Hamerstrom, 1940; traplines are always empty, e.g., dense vegeta- Sterling, 1953; Baker, 1983; Metzgar, 1967; tion subject to spring flooding (Table Rod-12). Voight and Glenn-Lewin, 1978). In southeastern Wisconsin, Popp et al. Parasites include trematodes, tapeworms, (1989) found no regular cycles, but following many species of nematodes, and one pentas- shortly after an ice storm, which knocked down tomid worm (Whitaker, 1968). Also found many branches adding to the debris on the infesting these mice are external mites, chig- forest floor, a peak of Peromyscus leucopus gers, ticks, sucking lice, and bot fly larvae resulted. Might this suggested cause-effect in- (Doran, 1954) as well as a few internal proto- deed have resulted from the storm? Can cli- zoans. The bot warbles fill the groin area quite mate generally, or a severe winter or deep frost, often, until the larvae emerge from the hosts, or an ice storm, affect density dramatically, in but to not sterilize the infested mice (Miller a species considered non-cyclic? On the prai- and Getz, 1969; Timm and Cook, 1979; ries of the Buena Vista Marsh, where meadow Timm and Lee, 1982). Whitaker (1968) lists voles and some P. m. bairdii were taken over internal parasites for P. leucopus, but he does 25 years of study (Hamerstrom’s journals), not record where the infested and infected there were surprisingly few leucopus caught mice were collected. (Nora Lopez-Rivera, personal corr.). One Home Range and Density. The classic “peak” (judging from recorded measurements) work on home range and territoriality in small was actually numbers of two species of deer mammals was written by W. H. Burt (1940) mice (all formerly considered as leucopus), and was based in part on deer mice studied in when identified from the records. nearby lower Michigan. Burt found that pop- Whether or not mast cycles relate to cy- ulations range from about four per acre in clic abundance in P. leucopus is obscured in spring to 12 per acre in the autumn. Over a Wisconsin by the ubiquitous habitat selection 23-year trapping period in central Illinois, of the mouse (see Habitat). The rodent in- trapping for 3-day periods in late summer and habits and seems to prefer oak stands, even autumn, produced one to 39 mice per acre flourishing near scattered oaks in cut-over (Hoffmeister, 1989). Half the population was maple-birch hemlock forest. This mouse subadult in winter and summer (Batzli, 1977). hoards acorns for winter. In oak forests in In upland woods Batzli found as many as 16 / southeast Wisconsin and in savannas in cen- ha in November, but as low as 3/ ha in Sep- tral Wisconsin, the numbers of white-footed tember. Even higher densities than those mice increase in mast peak years and decrease above were found by Blem and Blem (1975). when acorns become scarce, as observed in In central Wisconsin (in mostly jack pine- eastern oak forests (Ostfeld et al., 1996). oak savanna and shrubby, sandy prairies, and a Acorn abundance “controls” densities of P. few white pine forests) surveys based on tra- leucopus, P. maniculatus, and Tamias stri- plines at 10 m intervals and with 2 traps per atus in eastern forests. station, usually in transects 40 to 100 m in Inasmuch as the white-footed mouse is length, Long (1973) found absolutely no mice such an able climber, its home range has a in line after line following a hard winter (with vertical dimension (Horner, 1954). With a deep frost). Only in a few ravines, one taken flashlight I have seen them perched 12 feet near springs of water, were carry-over adults up small trees, and had them winter-nest in trapped. Evidently some survived elsewhere, my porch siding above the windows (in a nest because by June young were appearing in trap made of rabbit fur). Gosling (1977) found half

TAXONOMIC ACCOUNTS / ORDER RODENTIA 249 the mice caught living in nests in trees as high footed mice live in different habitats and vary as 13 feet. In competition with our long-tailed slightly in form adapting to them. (P. leuco- Peromyscus maniculatus maniculatus, the pus from a mesic hemlock habitat will be bet- white-footed mouse tends to occupy lower ter furred, darker brown, and longer tailed, branches and nest sites. Burt’s (1940) esti- whereas in a sandy dune area the same spe- mate of home range (ground area only) was cies may be small, scantily haired on the tail, about 0.27 acres for males and only 0.21 and surprisingly even slightly smaller of foot.) acres for females. I follow Osgood (1909) in referring all white- Analysis of the geographic distribution footed mice in Wisconsin and most of Illinois, in Door County and northern Wisconsin sug- for that matter, to the race P. leucopus noveb- gests that ecological replacement has occurred oracensis for the reasons detailed above. (Long, 1996). Long also found that an inva- Specimens examined. Total, 897. This sion of Washington Island, in Lake Michigan, is the largest sample of Wisconsin mammals of P. leucopus was coincidental with a sharp in the UW-SP mammal collection. Adams, reduction in the incidence of P. manicula- Ashland, Bayfield, Burnett, Chippewa, Co- tus, but maniculatus still persisted in the lumbia, Crawford, Dane, Dodge, Door, wetter swamps and forests. Now P. leuco- Douglas, Dunn, Eau Claire, Fond du Lac, pus has suffered something of a drop after Forest, Grant, Iowa, Jackson, Jefferson, becoming widespread and abundant. Both Juneau, Kenosha, Kewaunee, La Crosse, species seem to be fluctuating in density and Lafayette, Langlade, Lincoln, Manitowoc, the winters are continuing mild (see Fig.). Marathón, Marinette, Marquette, Milwau- (Hypothetically, the white-footed mouse kee, Monroe, Oconto, Oneida, Outagamie, seems less adapted to cold winters.) P. leuco- Ozaukee, Pepin, Pierce, Portage, Racine, pus is now established in hayfields, hedges, Richland, Rock, Rusk, Sauk, Sawyer, Sha- on the beach sand dunes, roadsides, and of wano, Sheboygan, St. Croix, Taylor, Trem- course in the forests. If ecological replacement pealeau, Vernon, Vilas, Washburn, Wash- occurs it seems likely slow. In 1991, the rela- ington, Waukesha, Waupaca, Waushara, tive numbers indicated that replacement might Winnebago, Wood counties. occur rapidly. Remark. Dental abnormalities in deer mice are rare, but Zielinski (1978) described a Peromyscus maniculatus (Wagner) small, supernumerary and anomalous tooth Forest Deer Mouse erupting labially, next to the upper first molar. Additional Natural History. Lackey et al. “The internal cheek pouches enable the mouse to (1985) reviewed natural history for the white- gather and carry small seeds to its secret grana- footed mouse, P. leucopus. ries... in holes of trees or under logs and stumps.” Geographic variation. According to — W. H. Burt, The Mammals of Michigan Hoffmeister (1989) there is a cline in size in P. leucopus so that some western popula- The taxonomic problem with the long- tions are similar to the nominate race. He tailed mice was discussed early by Coues and found no significant geographic variation in Allen (1877) and authoritatively by Osgood Illinois. I found none in Wisconsin, and sus- (1909). The racial differences are subtle and pect the Wisconsin stock invaded in the Ho- confused by clinal variation. Osgood changed locene (i.e., Recent) epoch, from Illinois. existing classifications, including his own, by Therefore, in such a short time, geographic resurrecting the available name P. gracilis (Le variation in Wisconsin would be expected to Conte) and placing in synonymy the names be insignificant. From place to place, white- umbrinus Miller and canadensis Miller. P.

250 THE WILD MAMMALS OF WISCONSIN m. gracilis is a problematical name for two then the following arrangements might ap- reasons. First, the type and topotype used in ply, using the schematic map. (A) is the nom- the description and subsequent comparisons inate race, (B) represents mice from southern are not well made. Miller, an astute and care- Canada northward of Lake Superior, (C) re- ful taxonomist, had ruled gracilis out as any fers to Wisconsin mice, (D) refers to mice in available name, because the type specimen Lower Michigan with long tails. was so poorly made. He wrote that the tail, 1. ABCD may be combined together with although it appeared to be too long for P. P. m. nubiterrae (E) a distinct race. Then leucopus, had no pencil (or tuft), implying it ABCD would be a variable nominate race was P. leucopus. Coues and Allen also said it known as P. maniculatus maniculatus. was impossible to separate the two specimens Chiefly it would vary in tail length, a trivial from leucopus. But in those days no one quite character, and tone of upper pelage (yellow- knew what was P. leucopus and what was P. ish, dusky, brown). maniculatus. Osgood thought the long tail 2. If gracilis was valid, its holotype tru- distinguished the type from P. leucopus. Drs. ly from lower Michigan, the name may be used Al Gardner and Michael Carleton recently for the small, dark, long-tailed mice there. wrote me saying they examined the holotype Then ABC would be the nominate race and and concurred that the specimen was refer- nubiterrae probably a valid race. able to P. maniculatus. The type locality is 3. The name P. canadensis may be raised also questionable, being referred to three from synonymy and applied to B + C. Some states, and as for Michigan, it might have been insular populations (Isle Royale, Rock, St. on the Lower Peninsula in the range of P. Martin, possibly Washington Island) southward maniculatus. of these populations are referable (identical) to The arguments and prestige of Osgood’s the specimens I have examined from Labra- (1909) “before its time” monograph seem to dor’s maniculatus (A). Such action solves noth- have led to general acceptance of the name ing and describes variation poorly. gracilis. But in some places, Wisconsin and The current classification lumping BCD Minnesota for example, the size of the sam- together works geographically, but includes ples used was too small to provide such con- B with C when some of them are A, and in- crete taxonomy (Long and Long, 1993). A large sample from Isle Royale was referred by Osgood not to the mice known as gracilis immediately southward, nor really to those north of Lake Superior (formerly known as P. canadensis) which differed slightly from them, but to the northern race from Labra- dor. Southward the trivial character of average length of the tail (of 7 mice from two localities) was used to refer all Wisconsin mice to P. m. gracilis. Long and Long (1993) found some Wisconsin mice on islands were identical with series of Labrador P. m. maniculatus, some on the mainland were slightly different, and those from lower Michigan were slightly different from all the Wisconsin mice. If one considers P. m. nubiterrae to the Schematic map showing P. maniculatus populations eastward as a distinctive but similar race (E), A-E. See text.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 251 cludes D also which seems slightly smaller and are slightly longer than in P. leucopus. The darker than any of the others. Some on Lake total length of the skull averages longer than Michigan isles in C are identical to A. in P. leucopus, although some specimens The most practical assignment is to com- from the southernmost parts of the geograph- bine ABCD together as a variable race, rele- ic range are no longer. The elongate tail is gating the name P. m. gracilis (based on dubi- well furred, so that the color is always distinc- ous type) to synonymy. The small dark, Lower tively bicolored (brown above, white or buffy Michigan populations are only slightly differ- white below) and it is tufted at the tip. The entiated from mice in southern Canada, north- whiskers are longer than in P. leucopus cor- ern Wisconsin, and upper Michigan, but may related with the prominent rostrum (snout), retain the name P. m. gracilis thereby em- and lower ones are much longer and usually phasizing only a little geographic color varia- white). The ears in many specimens of P. m. tion. All the mice are morphologically similar. maniculatus are enormous, but some have ears no longer than in P. leucopus. General- ly this trait distinguishes the two, as does tail Peromyscus maniculatus maniculatus length. The combination (Long and Long, (Wagner) 1993) of ear length, tail length, skull length, Forest Deer Mouse and length of incisive foramina serve to distinguish at least 95 percent of the specimens. 1845. Hesperomys maniculatus Wagner. Arch. From the tiny, sand-dwelling P. maniculatus Naturgeschichte, Jahrb. 1 1 (1): 148. Type bairdii, with which the long-tailed forest mice from the Moravian settlements scattered along are reportedly conspecific (perhaps linked by the coast of Labrador. limited intergradation in a few western local- 1855. Hesperomys gracilis Le Conte. Proc. Acad. ities), there may be in nature some intergra- Nat. Sci. Philadelphia, 7: 442. Type possibly dation. There is no evidence of intergrada- from “Michigan”. tion in Wisconsin between P. m. manicula- 1893. Sitomys americanus canadensis Miller. Proc. tus and P. m. bairdii. Biol. Soc.Washington, 8: 55. Type from Pe- There are six mammae (4 pectoral, 2 terboro, New York. inguinal), although only four functional teats 1898. Peromyscus maniculatus: Bangs. Amer. Nat., are seen in some lactating females. The bac- 32: 496. ulum is a short, slightly curved rod. John Long 1897. Peromyscus maniculatus umbrinus Miller. and William LeGrande (unpublished) deter- Proc. Boston Soc. Nat. Hist., 28: 23. Type mined that the number of chromosomes is from Peninsula Harbor, north shore Lake Su- 48, both in northern Wisconsin and on Wash- perior, Ontario. ington Island. See Singh et al. (1966). 1908. Peromyscus canadensis (Miller): Jackson’s A There is no great difference between Preliminary List of Wisconsin’s Mammals. Bull. Wisconsin and nearby Upper Michigan mice Wisconsin Nat. Hist. Soc., 6(1-2): 13-34. from those on Lake Michigan isles. In the Beaver Islands there is insular variation with The name Peromyscus means little some populations resembling those of lower mouse, but the prefix Pero has no certain Michigan. The specimens that I have studied meaning. The specific part of the binomen, from there are slightly smaller and darker maniculatus, means small-handed. brown. Essentially there is a little difference Description. The skull is smoothly round- in them and any Wisconsin mice. These also ed with elongate rostrum and elongate pala- resemble the Isle Royale mice, which have tal slits (incisive foramina). These slits are near- been ascribed by Osgood (1909), himself, to ly parallel, i.e., seldom pinched anteriorly, and P. m. maniculatus. In most of Wisconsin

252 THE WILD MAMMALS OF WISCONSIN there are some mice with a yellowish tone, see account of that species and Long and less brown than in Michigan. Often the Wis- Long (1996). consin mice are russet-brown or a dusky Status At this time the populations of P. brown. There is no difference apparent from m. maniculatus are not in any serious peril, them and the Labrador mice I have exam- and they are beneficial mammals because they ined. The venter is whitish, sometimes help control forest insects, and are prey for creamy. In a small population on the east side forest raptors and predators. They probably of Washington Island, Ron Zimmermann and help plant forests because they scatter-hoard I once collected a series with some venters acorns, beech nuts and other seeds. William tinged with pink or pink-ochraceous. This is Henry Burt, probably the most famous mam- a curious but probably transient variation. malogist in the history of Michigan, called this Jackson (1961) reported that on the Upper attractive, large-eyed mouse the “most beau- Peninsula of Michigan these deer mice often tiful mammal in Michigan.” showed white spotting (possibly from para- Habitat. This nocturnal long-tailed climber site bites?). The ears are brownish gray often is found in beech-maple forest with hemlock, edged with a thin margin of white, and the cedar, yellow birch, and some oak. The mice eyes are a shiny black. The feet are whitish, also live in swampy, boggy, and riparian thick- as is the underside of the tail, and in this spe- ets in spruce-fir, maple, pine, or mixed forests. cies the dark upper side of the tail may show They inhabit dune heath, in Upper Michigan, only as a slender brown line with much of the but not to my knowledge in Wisconsin. Often white showing, even in dorsal view. The pen- the soils are rocky, with outcrops and ravines. cillate tuft is a mixture of brown and white The Niagra escarpment provides many bluffs hairs. There is an intermixture of black hairs and cliffs for this mouse on Lake Michigan isles. dorsally, but not so evident as in P. m. bair- More detail has been compiled on habitats by dii. The juveniles are gray above and white Hooper (1942) and Long (1996). below. They are difficult to distinguish from The long-tailed P. m. maniculatus is P. leucopus (except the ears in P. leucopus more adapted to winter conditions than is P. are shorter). Adults molt annually in summer leucopus, including the tendency to enter (Osgood, 1909: 19-21). Some adult males are torpor, build larger nests, make larger food about 5 percent larger than adult females. hoards, diminish food consumption, show Dental formula. I 1/1, C 0/0, P 0/0, M regression of sex organs, fur out the pelage, 3/3 = 16. and so forth (Wolff, 1986; Wolff and Hurl- Geographic Distribution. The nominate butt, 1980; Vogt, 1993). race is found primarily in the forests of northern Wisconsin and Lake Superior, and most of the islands in Wisconsin waters of Green Bay and Lake Michigan. This species does not occur on the Door Peninsula. Its southern boundary extends somewhat northward of Green Bay across the state, to the Rib Riv- er near Athens in Marathon County, and northwest to and beyond the state boundary of Minnesota. There are two relict populations south of the boundary, but the Sheboyg- an Marsh mice seem eradicated. See Map. For additional evidence regarding ecological Skull of Peromyscus maniculatus maniculatus. Note replacement of this species by P. leucopus, elongate palatal foramina (i).

TAXONOMIC ACCOUNTS / ORDER RODENTIA 253 These deer mice nest in tree hollows, examined. Some stomach contents seemed stumps, and possibly abandoned birdnests. berry stained, and seeds became more preva- Such a scansorial species tends to nest high- lent in summer. Small arthropods comprise er in trees and seldom does so on the ground part of the diet. In some stomachs the eggs (in comparison to P. leucopus) (Horner, and jaws of carpenter ants were observed. Jack- 1954; Stah, 1978; 1980; Wolff and Durr, son (1961) lists as foods, beaked hazelnut, 1986: Wolff and Hurlbutt, 1982). Only three Juneberry (Amelanchier), basswood, winter- of 39 P. maniculatus nests were underground green, pin cherry, hemlock, maple, blueberry, whereas 36 were in trees. Nest heights were yellow Clintonia, and partridge berry. On Lake significantly higher (7.4 m to 4.3 m) in P. m. Michigan isles Hatt et al. (1948) found evidence maniculatus than in P. leucopus. at the nests that this species scavenges on cray- The long-tailed P. maniculatus some- fish, small fish, even bird bones. Some remains times invade houses to gather fibrous materi- were cached. This mouse has large internal al for a nest. On Washington Island a friend cheek pouches (Burt, 1940) to transport foods told me a mouse had transported rug fibers for hoarding. There seems no marked differ- in excess of 100 m to its nest. Usually the ence in the diets of this species and P. leuco- nest is a wad of grass stems, mixed with leaves pus (Zegers and Merritt, 1988). and bark (Cory, 1912). On Lake Michigan Reproduction. Little information is isles the nest materials included moss, bird known of reproduction in Wisconsin for this feathers, milkweed down, and dry vegetation race. The mice commence breeding in March (personal observations). (Jackson, 1961, suggests late April), and con- Foods. A few grass seeds, acorns, old tinues into September. The litter size is re- beech nuts, and several arthropods were all I ported as 3-5. Possibly four litters are brought could find available for deer mice to eat in June forth in a single growing season by one mother on Washington Island. Stomachs seemed to (Burt, 1940), but if the pattern resembles that contain these foods finely minced. Possibly in other Peromyscus, which seems likely, then these mice eat green leaves then (Clark, 1972); many young-of-the-year may breed once or I found no green materials in the stomachs twice in summer and fall, and after winter passes those that survive it may breed once or twice again the following spring. The UW- Table Rod-12Rod-12. Percent of Washington Island Peromy- SP museum collection has few specimens of scus found in three habitats. Long, 1996. pregnant females, and Jackson (1961) could Sand Old Beech-Maple find no young in July (1919) on Outer Island, Dunes Fields Forest in the Apostles. In other Wisconsin Peromy- 1975 scus, the young enter the populations in June. P. leucopus 00 0On Washington Island I caught young in ear- P. maniculatus 0 0 100 ly July. From a litter of five weaned mice, I kept one six years in the laboratory and an- 1984-1986 other one almost as long (and subsequently P. leucopus 00 0 P. maniculatus 0 0 100 may perish, although they are long-lived). Mortality. Because this mouse is difficult 1987-1990 to identify in scats or owl pellets, little is known P. leucopus 10 5 55 of its predators. P. leucopus may prey on the P. maniculatus 00 29young and may drive them from their homes 1991-1994 (see account of P. leucopus). When present, P. leucopus 21 11 50 Blarina surely preys on these mice. Snakes, P. maniculatus 00 17owls, and other birds of prey doubtless take

254 THE WILD MAMMALS OF WISCONSIN them. Raccoons rob my traplines of mice on Washington Island, and doubtless eat the young if they can get them in trees or on the ground. Weasels, martens, fishers, foxes and other carnivores eat Peromyscus whenever possible. Forest fires destroy them, and tree-cutting, logging, and natural succession remove habitats, which reduces the populations. Two kinds of lice, tick, flea, and mite infest this mouse according to Jackson (1961) who mentions botflies and internal worms. I observed botflies on mice from Washington

Map showing geographic distribution of Peromyscus maniculatus maniculatus in Wisconsin and for P. maniculatus in North America. See account of P. m. bairdii.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 255 Island. P. m. maniculatus usually seem clean ing from Labrador in eastern Canada west- of mites and fleas. ward through Ontario and into Minnesota, Home Range and Density. In northern and extending southward into northern Wis- Michigan, Manville (1949) found the home consin, Upper and Lower Michigan (persist- range to vary between 500 to 1,500 square ing also on most of the islands and even small yards, a fraction of an acre. Juveniles had smaller isles in Lake Superior and Lake Michigan), home ranges. Blair (1942) determined the home the long-tailed Peromyscus maniculatus are range to be more than 5.5 acres in males, 3.75 morphologically similar. Mice in lower Mich- acres in females. See table Rod-13. igan differ from the others in uniform brown Breeding females were seldom caught upper parts and small size, with the tail actu- near one another, but males were frequently, ally or relatively a bit longer. If mice in south- indicating the females are territorial. (Or they ern Canada, Upper Michigan, and Wisconsin may stay near their nests.) The home range differ from the typical mice of Labrador, it is has vertical dimension that has not been stud- in slightly smaller size of skull, especially the ied On foggy nights the mice may be seen rostrum, and in a more often yellowish brown climbing in trees. color. The tail tends to longer size. The typi- Occasionally a winter cabin in the North- cal mice of the nominate race have dark up- woods is overrun with these mice. One autumn per parts, a shorter tail, and robust rostrum and early winter on Rock Island, Tom Jessen and skull. These traits are seen in mice on caught nearly 100 in one of the “Icelandic” Isle Royale, and of mice from the Islands of houses there. Jackson (1961) reported from the Green Bay and some other isles in western field notes of Clarence Birdseye, of subsequent Lake Michigan. Washington Island mice were frozen foods fame, who worked for the U. S. slightly smaller than those on Rock, St. Mar- Biological Survey, that in Luce County, Michi- tin, and Big Summer islands. gan, in the summer of 1910, he caught a total On the Beaver Islands across the Lake in of 61 mice in one cabin. He saw 8 drowned in eastern waters, the mice are somewhat small- a kettle, and dogs, cats, and poisons killed many er and darker brown, at least on some of the more in surrounding shacks. On Washington islands there, but they average a little larger Island I counted as many as 45 to the acre in than those I examined from the Lower Penin- beech-maple-hemlock forest. In the fields and sula. In northern Wisconsin there is fluctuat- along rock walls I caught none. Usually several ing variation in both size and color of exam- are taken in each trapline in suitable habitat on ined specimens, with narrow rostra and yel- all the Green Bay islands (except Poverty Island lowish pelage tones prevailing. Specimens from which is dominated by Clethrionomys). In the Ontario are indistinguishable from them; and northern forests they were never reported to that resemblance of Ontario specimens to P. be as abundant as observed densities in peak m. maniculatus was mentioned by Osgood years on Washington Island. (1909). He felt the mice on Isle Royale (a large Remarks. Not only are these mice beau- sample) were typical of P. m. maniculatus. In tiful; they are gentle and docile, not leaping the Apostle Islands the deer mice are variable, away as does P. leucopus or biting the fin- from isle to isle, but not so closely resembling gers. They are cautious climbers, and one can the Labrador specimens. In southern Wiscon- hold them (but do not alarm them) so long as sin, the relict mice are small and dark perhaps they are not allowed to climb your arm or resembling the mice in Lower Michigan. Prob- escape on one’s desk. ably many years ago the small, dark brown Geographic variation. Although there are mice were distributed continuously around the subtle differences from one population of south end of Lake Michigan. I refer the forest these long-tailed deer mice to the next, rang- deer mice all to the nominate race.

256 THE WILD MAMMALS OF WISCONSIN Table Rod-13. Home range in acres in adult deer mice in scription seem more apt to describe a speci- nearby Michigan. After Stickel, 1968. Based upon work by men of Mus musculus (the number of mam- Blair and Manville. mae, the position and protuberance of the eyes Species and Sex Habitat/ Home Range and so on). Indeed, the authors themselves use the name Mus. Other authors (see below) did P. m. maniculatus Conifer-Deciduous Females June:0.19 apply this name to the prairie deer mouse, and September: 0.1-0.31 not to P. leucopus, but their publications’ dates Males June: 0.18 are later than that of the name Mus bairdii September: 0.12-0.25 Hoy and Kennicott. P. m. bairdii Beech Maple 1846. Mus Michiganensis Audubon and Bachman. Females August: 2.31 September: 0.88-5.64 Viviparous Quadrupeds of North America. Males August: 1.39 Same description verbatim from the earlier September: 0.79-3.28 description, applied the same way. P. leucopus Bluegrass 1857. Mus bairdii Hoy and Kennicott. Agric. Report Females August: 0.63 U. S. Commissioner of Patents, 1856. Page 92. September: 0.25-1.67 Males August: 0.51 Type from Bloomington, McLean Co., Illinois. September: 0.12-2.29 1883. Hesperomys Michiganensis (Audubon and P. leucopus Oak-Hickory Bachman). In Moses Strong’s List of mammals Females August: 0.27 in Wisconsin. Geol. Surv. 1873-1879. Page 439. September: 0.16-0.54 1902. Peromyscus bairdii: Snyder. Bull. Wisconsin Males August: 0.21 September: 0.006-0.37 Nat. Hist. Soc., 11, p. 116. 1908. Peromyscus michiganensis (Audubon and Bachman): In Jackson’s A preliminary list of Specimens examined. Total. 309, and Wisconsin’s mammals. Bull. Wisconsin Nat. from Michigan 52. Ashland, Door, Douglas, Hist. Soc., 6(1-2): 13-34. Florence, Forest, Iron, Langlade, Lincoln, 1909. Peromyscus maniculatus bairdii: Osgood. N. Manitowoc, Marathon, Oneida, Price, Saw- Amer. Fauna, 28: 79. yer, Sheboygan, Taylor, Vilas counties. Other records. Ashland Co.: (Kantak, The name Peromyscus means little 1981): Madeline, Oak and Sand isles. mouse although the prefix Pero is unknown. Michigan Delta, Charlevoix counties. The name bairdii honors the great mammalogist and museum administrator at the Smith- sonian Institution, Spencer Fullerton Baird. Prairie Deer Mouse The name maniculatus means small-handed Peromyscus maniculatus bairdii or small-footed. (Hoy and Kennicott) Some mammalian subspecies are so weakly differentiated that this one, which acts almost 1842. Mus Michiganensis Audubon and Bachman. everywhere as a “good” species, seems a taxo- J. Acad. Nat. Sci. Philadelphia, 8: 304. Type nomic mistake. If the “mistake” is not in the from Erie Co., Michigan (probably Ohio, ac- fact of supposed, indirect, and far-removed in- cording to Osgood, 1909). Osgood uses this tergradation, it may be in an arbitrary percep- name to describe a white-footed mouse, possi- tion of the importance of hybridization, local bly because of its long tail, and he placed the and limited versus wide-spread intergradation. name in the synonymy of P. leucopus noveb- There is some hybridization that takes oracensis. The same can be said for the name place in the laboratory (Foster, 1959; McIn- combination that follows (Audubon and Bach- tosh, 1956; Dice, 1968), although fertility man, 1846). In my opinion the name and de- seems limited in hybrids. There is no known

TAXONOMIC ACCOUNTS / ORDER RODENTIA 257 hybridization eastward (with P. m. nubiter- eyes are large but not so large in relation to rae). Westward it becomes paler and may in- the snout as in Mus. The incisors are not tergrade with short-tailed pale mice, which grooved as in Reithrodontomys. There are merely may have higher frequencies of pale- 48 chromosomes (Singh et al., 1966), and ness genes. The pale mice do not seem to six mammae (four pectoral and two inguinal). intergrade much if any with neighboring long- The baculum is a slightly curved rod. tailed forms. Therefore, P. m. bairdii and P. Dental Formula. I 1/1, C 0/0, P 0/0, M m. luteus of the Great Plains seem conspe- 3/3 = 16. cific. Osgood (1909) thought that in the south- Geographic Distribution. This little deer western part of its vast geographic range P. mouse is fairly common in suitable sandy and m. bairdii might intergrade with western long- prairie habitats in the southern parts of Wis- tailed mice. This western area should be the consin (see Map). It has extended its geographic focus of taxonomical studies, to resolve a range northward into Upper Michigan in those problem for deer mice important in at least beach dunes, mostly fringing Lake Michigan’s 11 states and one Canadian province. western shore, and prairie areas near Mari- Description. The adult fur is almost rus- nette. There is a relict population isolated on set or chestnut brown above and creamy be- the dunes of the Door Peninsula, where the low. There is a strong intermixture of dark hairs other (long-tailed) Peromyscus maniculatus is intense dorsally along the mid-line. The gen- unknown (Jackson, 1961). Recent collections eral tone is dark brown. The upper side of the from the dunes at Algoma and as far north as tail is nearly black, the underside white. Occa- Northport yielded the first specimens known sionally the adults are grayish or tawny brown to me of P. m. bairdii from the Door Peninsu- instead of dark or chestnut-brown. Young mice la in over 60 years. Two UW-Madison speci- are juvenal gray above, white below. mens obtained by Frank Iwen, Univ. Wiscon- The meaning of the latinized name, sin–Madison, were taken near Algoma. “small-handed” actually refers, of course, to Clearing of forests and creating railroad the forefeet, and is truly relevant for identifi- and highway rights of way encouraged the cation. Only harvest mice and house mice (see mouse to disperse northward. This mouse accounts of those species for their character- occurs in southern counties on the opposite istics) have feet as small. The hind foot is al- shore of Lake Michigan, lower Michigan most always 17 or 18 mm in length. When (Hooper, 1942). on occasion it reaches 19 mm it becomes dif- Status. This mouse seems to do no harm ficult to distinguish the mouse from P. leuco- to humankind. It dwells in sparse grasses in pus. The foot is so obviously slender and delicate (with tiny toes) that recorded measurements (which may be in error) may be misleading. The short (up to 64 mm in length), stout tail is distinctive. In young white-footed mice the hind foot is usually at least 20 mm in length and the tail is attenuate and slender (Long and Long, 1993). The total length is less than 160 mm. See Key. The skull is smooth and rounded, delicate in appearance. It never exceeds 24.5 mm in greatest length. The prominent ears measure less than 14 mm So-called races of Peromyscus in Wisconsin. Above P. in length. The ears are similar in shape but m. maniculatus, below P. m. bairdii. Note its short tail, ears, smaller than those in P. leucopus. The black and whiskers.

258 THE WILD MAMMALS OF WISCONSIN prairies or barren sandy fields and dunes. The even plowed grain fields. They are often found isolated population in Door County merits with harvest mice under shocks or bales of protection. hay, in thin stubble of well-drained pasture or Even if it is not a distinctive taxon, it may drained marshland, in weedy areas, railroad prove interesting to study. It is a semi-isolat- rights-of-way where Equisetum, grasses and ed population with narrow habitat preference. forbs are common, dwelling on the ground The specimens seem stunted, i.e., small, and beneath vines, hazelnuts, and small trees, on closely resemble P. leucopus, which in that sand dunes and sandy lake beaches, and oc- habitat also seems rather small. curring with P. leucopus when the brambles Habitat. The prairie deer mouse inhab- or brush is sufficient for the latter. its open fields and grasslands, without much The nest is never built in trees as seen in tree cover or brush, and well drained sandy other Peromyscus (those with long tails); this soils and thin soils on pastures, hayfields, and species builds it on the ground or at the end

Map showing geographic distribution of Peromyscus maniculatus baidii in Wisconsin. See map for P. maniculatus maniculatus for North American range. Line shows recent northern localities.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 259 of a short tunnel (Howard, 1949). The nest Succession of shrubs and trees, plowing by sited under a board, rock, shock of grain, or farmers, and fire probably destroy many. some other sort of rain protection, is about Parasites include some of the ectopara- 10 inches ( = 254 mm) in diameter, construct- sites of the other Peromyscus, and some of ed of grass or weed stems. Inside it is lined their endoparasites as well. Whitaker (1968) with thistle or milkweed down, fur or feath- lists internal and external parasites for Indiana. ers, or fine grasses. There is usually a single Home Range and Density. Home range is entrance. The nest is not re-used after once less than an acre (Burt, 1948). Males have slight- abandoned; a new nest is constructed. Hout- ly larger home ranges than females. Breeding cooper (1917) found burrows 16 feet long, females do not show much overlap in home and about a foot below ground. range indicating that they may be territorial. In Foods. Not much is known about the diet winter the prairie deer mice may aggregate to- in Wisconsin. It prefers seeds, fruits, and oth- gether in groups as large as 15 members. After er vegetal foods. Plum pits were observed in winter the deer mice disperse from their winter the nest or near it, and were observed in one quarters to commence breeding. winter nest. In barren soils density of P. m. bairdii Reproduction. Breeding begins in early may seldom exceed 9 per acre. Mice in na- March and continues into November. A fe- ture usually live no longer than 2 years, and male may bear as many as three litters each the turnover is considerable. Howard (1949a) year. The male is driven from the nest after found that of 1,200 mice less than 240 the young are born. Gestation is about 25 reached sexual maturity, and only 8 lived long- days. The litter size averages about 3-5 young er than 73 weeks. Seldom would one find a (range 2-9). Gestation is 23-27 days. The sharp peak of abundance. Nora Lopez-Riv- blind, naked young are rapidly furred out and era (personal correspondence) found a sharp two weeks the eyes open. Weaning takes place peak in density in the records of Fred and after 25 days of post-natal care (22-37 days Fran Hamerstrom, kept faithfully over a peri- according to Svihla, 1932). od of 25 years. Analysis of the hind foot and Apparently P. m. bairdii has the same tail measurements carefully recorded revealed pattern of breeding as does P. leucopus. that some of the high-density mice were in Winter carry-over mice breed in spring. There fact P. leucopus. Therefore, there was no is a pause in breeding in mid-summer and peak for P. m. bairdii. They are seldom tak- reproduction recommences in late summer en in high numbers in the prairies of the Bue- and autumn. Litters are brought forth as late na Vista Marsh or the sandy hazelnut and as November. savanna areas of central Wisconsin, but in Mortality. Hardly anything is known of southern Wisconsin they were taken in fair predation on this species in Wisconsin. It is numbers by Stromburg (1979). difficult to obtain enough remains from fecal Geographic Variation. None has been scats and bird pellets to identify this species observed in Wisconsin. from the other deer mice. Jackson (1961) lists Specimens examined. Total, 150. Ad- known predators as striped skunk, weasels, ams, Bayfield, Buffalo, Burnett, Columbia, raccoon, red fox, and coyote. Domestic cats, Dane, Door, Dodge, Eau Claire, Fond du and on the Buena Vista Marsh in central Wis- Lac, Iron, Iowa, Kenosha, Kewaunee, La consin, and probably many other marshes, Crosse, Manitowoc, Marathon, Milwaukee, the harrier (= marsh hawk), and kestrel prey Monroe, Ozaukee, Pepin, Portage, Price, on these mice. Saw-whet, short-eared, long- Racine, Rock, Rusk, Sauk, St. Croix, Ver- eared, boreal, and great-horned owls eat these non, Waukesha, Waupaca, Waushara, Win- mice. Doubtless prairie snakes capture some. nebago, Wood counties.

260 THE WILD MAMMALS OF WISCONSIN 2 Molars with deep re-entrant angles on one side only, opposite side comprised of sinuous arcs; each upper incisor with small groove extending the length of the tooth ...... Synaptomys 3 3 Lower molars with shallow, sinuous re- entrant angles on outer (i.e., labial) side; 3 large and 1minute prisms on lower mid- Upper right tooth-rows of Wisconsin microtines. The dle molar ...... Southern bog lemming patterns are as distinctive as signatures on antique vases. Synaptomys cooperi Pitymys ochrogaster and P. pinetorum are identical, Microtus 3’ Lower molars with little indentation (re- pennsylvanicus (middle) has a distinctive posterior islet on entrant angles) on labial side; 3 large the middle molar. There is neat cusp alignment (Clethrionomys prisms on lower middle molar; known in far right) and deep re-entrant angles on one side only in Synaptomys (second from right. After Hazard. Wisconsin or Michigan only from fossils ...... S. borealis 2’ Molars with re-entrant angles on labial and lingual sides, upper incisors lacking Subfamily ARVICOLINAE Gray grooves ...... 4 (=Microtinae Schrank) 4 Teeth small with neatly regular indenta- Voles and Allies tions, sometimes more curved than pointed, having relatively thick enamel bor- The voles and bog lemmings are difficult to iden- der; teeth rooted in adults; posterior loop tify, and one often must resort to examination of third upper molar irregular in shape; of their skulls and teeth. See Fig. above The lower anteriormost loop deeply constrict- much larger muskrat with its laterally com- ed as in meadow vole, with only 3 closed pressed and scaly tail is well known. The small- prisms and a posterior loop ...... er microtines may be identified by use of the ...... Red-backed vole following key, which excludes the muskrat. Clethrionomys gapperi 4’ Teeth larger; posterior prism of middle upper molar terminating abruptly in a Key to the skulls of voles and bog shoulder or bulge confluent with posteri- lemmings. After Long, 1990 or salient angle; posteriormost loop of third upper molar spear-shaped (subtri- 1. Upper middle molar with anterior loop, angular); lower anterior molar slightly three closed prisms, and one distinctive, constricted anteriorly and deeply con- small posterior loop or islet; posterior stricted posteriorly, with 3 prisms and a upper molar with anterior loop, three posterior loop ...... Pitymys 5 closed prisms, and distinctive posterior 5 Tail exceptionally short; fur fleecy, wal- crescentic loop; anterior lower molar nut or reddish brown; forefeet with ro- deeply constricted with five closed prisms bust claws; skull broad, with interorbital and a posterior loop ...... Meadow vole breadth more than half the length be- Microtus pennsylvanicus. tween the tips of the nasals to the poste- 1’ Upper middle molar with only three rior ends of the premaxillaries ...... prisms; posterior upper molar with two; ...... Pine or Woodland vole posterior loop not crescentic; anterior Pitymys (= Microtus) pinetorum lower molar with fewer than five closed 5’ Tail somewhat short, fur coarse and griz- prisms beween loops ...... 2 zled, buff or orange-brown; forefeet

TAXONOMIC ACCOUNTS / ORDER RODENTIA 261 small; skull narrow, interorbital breadth anterior border of the pterygoid fossa (i.e., about half the length between the tips of posterior nares). The teeth are narrow and the nasal bones and the ends of the pre- small. The upper third molar has an anteri- maxillaries ...... Prairie vole or loop, three closed prisms, and a com- Pitymys (= Microtus) ochrogaster plex, irregular posterior loop, resembling the pattern in Microtus. The more arcuate Genus Clethrionomys Tilesius angles are more even in linear alignment, (=Myodes Pallas 1811, ?nomen oblitum) and much more delicate with the tiny prisms Red-backed Voles “neatly outlined”. The upper middle molar consists of an anterior loop and three closed The teeth are small and primitive, reportedly prisms (which is ordinary in arvicolines). The rooted in adults; salient angles less pointed (i.e., lower first molar might be said to terminate more arcuate) than in Microtus or Pitymys; with two posterior loops in tandem, only upper third molar elaborate and irregular in two enclosed areas intervene between the form; posterior hard palate shelf-like, a trans- complex anterior part and the last loop. The versely edged palate. The dorsal fur is reddish, anterior portion is pinched so that a tiny and the venter whitish or cinnamon buff. inner salient may occasionally be closed off I provisionally retain the well-established as well. The third lower molar is distinctive Clethrionomys (= Myodes by priority). See in its three similar and large outer salient above. angles all in a row, and three small inner salient angles neatly arranged opposite. There are four pairs of mammae in females, Clethrionomys gapperi (Vigors) two pectoral and two abdominal. Small flank Southern Red-backed Vole glands are often evident in some males. The baculum has three distal processes and a 1830. Arvicola gapperi Vigors. Zool. Journal, 5: basal stalk. There are 2N = 56 chromo- 204. Type from between Lake Simcoe and somes (Merritt, 1981). Toronto, Ontario. The red-backed vole is easily identified, 1928. Cleththrionomys gapperi gapperi: Green. J. except in juveniles, which are dark brown. The Mammal., 9: 255. dorsum is russet or chestnut reddish brown, set off by ochraceous (almost yellowish)-gray- The name Clethrionomys means alder ish-tan sides. The sides are usually flecked with mouse. The trivial part of the binomen honors a Dr. Gapper from Ontario. Description. This small red-backed vole has a lightly built skull. The prisms and loops in the cheek teeth are neatly aligned and the teeth are small. Reportedly the teeth are often rooted in adults. Ostensibly a primitive feature, though shared by Phenacomys, rooted molars provide the basis for this species being mentioned first among the Wis- consin arvicolines (formerly microtines). The incisors are usually pale yellow. The skull is small and rather circular in profile. No other arvicoline has such a shelf-like or straight pos- Partial skull of Clethrionomys gapperi. Note hard palate terior border of the palate (it demarcates the shelf and molar teeth.

262 THE WILD MAMMALS OF WISCONSIN dark blackish, a false pattern of guard hairs, but they are actually gray bases of the hairs showing through the pelage. A few brownish guard hairs in dorsal and lateral pelage are evident. The ventral fur is either a grayish or pure white, although up to 40 percent at some localities may show pale ochraceous or cinnamon buff. The pale coloration is conspicuous on the throat and lower cheeks, often extending as a ventral line to whitish feet and claws. The general effect is of a tricolored mouse, red-brown above, gray orange on the sides, and whitish below. The tail is medium

Maps showing geographic distribution of Clethrionomys gapperi in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 263 in length. Six tubercles are reported on each Logs and stumps strewn over the forest floor hind foot. The coloration in juveniles appears seem ideal for this vole. They occur in hard- as a rich rusty brown with just a tinge of rus- woods, conifers, mixed woodlands, tama- set. The venter in juveniles is brownish or gray racks, quaking aspens, or silver maples in with just a tinge of white. Young mice in win- wetlands, but are most common in cedar- ter attain adult coloration, which fur devel- beech-hemlock, pine-maple, tamarack-maple- ops rapidly, faster than in summer. Molts re- hemlock, and spruce-fir forests. Red-backed portedly are in autumn and spring. voles thrive where deep snow lingers into The coloration (reddish dorsum, whitish spring. Getz (1968) suggested they prefer venter) and dentition clearly distinguish this habitats near water. The species never oc- species from all other arvicolines in Wiscon- curs in grasslands, but may make short and sin. The longer tail clearly sets a red-backed poorly defined runways in ground vegetation vole apart from the short-tailed woodland (= near its home. It occasionally digs burrows pine) vole and southern bog lemming, which resembling the tunnels of the shrew Blarina, do resemble dark Clethrionomys. and it tunnels under deadfalls, logs, torn out The tail is short to medium, and the ears tree roots, rock outcrops (personal observa- rather prominent for voles. Jackson (1961: tions), and decaying stumps (see Fig.). 225) gives external measurements as follows: Two nests were described by Jackson Total length ranges up to 150 mm, tail only (1961) in Price County. These were excavat- 32-42 in adults, hind foot 18-20, ear 14-16. ed by following the tunnels. The first was Weights vary from 25 to 36 grams. Total found at the end of a side branch ten inches ( length of skull ranges only to 24.8 mm, with = 254 mm) in length, not off the main tun- the width of cranium varying from 12.0 to nel. It was surrounded by tree roots. The nest 13.6 mm. was a carpet of grass stems, parts of dead Dental Formula. I 1/1, C 0/0, P 0/0, M leaves, and mosses. The nest carpet entrance 3/3 = 16. of a tunnel led Jackson to find a second nest. Geographic Distribution. Woodlands and The main burrow led about 14 feet (4.3 m) swamps as far south as Dane County. This from one elm to another and thence to a rot- boreal mammal is found on islands in Lake ted stump under which (at a depth of 18 inch- Superior and Lake Michigan. The species does not occur in the deciduous woodlands and prairies of southern and western Wisconsin. See Map. Status. The red-backed vole is fairly common although seldom abundant, occurring in boreal communities and definitely beneficial to humankind. A valuable component of northern food chains, it eats many seeds and seldom comes in contact with human agriculture. Habitat. Red-backed voles are found in the boreal forests in Wisconsin, occasionally in swamps and marshes if some trees are present. They may be found on uplands and slopes of hills, or along sloughs and stream valleys. The soils may be rocky, of black dirt, Burrow system of Clethrionomys gapperi. After Jackson sandy loam, overlain with leaf litter, or brush. (1961) and C. A. Long’s notes.

264 THE WILD MAMMALS OF WISCONSIN es) the nest was found. The burrow extended home ranges (Mihok, 1979; Perrin 1981). about 3 inches below the surface. At each Body weights drop in winter, so often tiny tree there were several branch passages drop- voles in adult pelage are taken in traps. Most ping to depths of 12 to 18 inches. The nest reproduction ceases then, but voles may breed was under a buried root of the rotted stump. in winter. In our collection there are juveniles This nest was about 4 inches in diameter and taken in winter. Both lactating females and one inch deep, and comprised of a slightly juveniles were taken in late November. Most hollowed but uncovered clump of dry leaves, young enter the population in summer and bark, hazelnut shells, hemlock cones, twigs, September. There is a postpartum estrus. sphagnum, and green mosses. The mosses Svihla (1930) calculated the gestation period were uppermost in the nest. Svihla (1930) as about 17-19 days. Two litters 18 days apart reported grass-lined nests, and Banfield (1974) suggest weaning may take place in 17 days. found nests in Canada occasionally in tree There could be four litters per summer, but in cavities or in limb crotches as high as 20 feet the usual Wisconsin pattern carry-overs usu- above ground. Nests can even be sited wit ally produce spring young, and young-of-the- hin tunnels in the snow. year produce autumn young. Probably Foods. The food contents in the stom- Clethrionomys follows this pattern. They may ach of the red-backed vole are finely ground. not produce four litters in a year, but several In Wisconsin they have been seldom exam- over 12 months. Some voles live up to three ined. Red-backed voles opportunistically roam years (Banfield, 1974). the woods feeding on plant parts, berries, and In Wisconsin the mean litter size was 4.75, nuts. During the growing season they feed with two modes (3, 6), and I observed a range on grasses and in summer the contents of the of 3-6. The reported range is 2-8 for the spe- stomach are often stained green. Hamilton cies. Only seven pregnant females were in this (1941), in New York, found that nearly three collection, taken from February to late Septem- fourths of the food was green vegetation finely ber (except for a few winter breeding records). minced. About 10 percent of stomach con- A female from Poverty Island, Lake Michigan, tents were remains of insects. Some spiders, in Michigan, had six embryos in August. snails and centipedes were also eaten. Fungi, Svihla (1930) described the young born bark and roots supplement the summer food naked with eyes closed, each weighing about (Schloyer, 1977; Banfield 1974). In autumn 2 g. Fur begins to develop four days after birth. and winter the seeds of beech, pine, cherry, The incisors erupt by seven days, and the eyes alder and dogwood are eaten as are hazelnuts, open in 10 or 11 days. bark, buds, and grass seeds (Krefting and Ahl- Mortality. The predators of red-backed gren, 1974; Abbot and Quink 1970; Vickery voles include both hawks and owls because 1979). Jackson (1961) adds to this list seeds these voles are active day or night. Red-shoul- of maples, hemlock, spruce, pin cherry, shad- dered and rough-legged hawks, and an occa- bush, black alder, mountain ash, blueberry, sional red-tailed hawk catch them by day, and partridge berry, and Clintonia. Caches of nuts barred, great-horned, and screech owls take and seed were found including Rubus, them by night (Banfield, 1974; Errington et Amelanchier and Vaccinium, but the vole may al., 1940; de la Perriere 1970; and Jackson carry a food item into its burrow to eat it in 1961). James Fitzpatrick (field notes, 1964) security, giving the impression of hoarding observed a snowy owl hunting where Clethri- food. There are no known winter caches. onomys was the dominant rodent. The rare Reproduction. This vole is not sociable great gray owl occasionally takes one. Bears, (Manville, 1949; Merritt, 1981). Females raccoons, coyotes, wolves, red foxes, wea- seem to exhibit territorial defense of their sels, striped skunks, badgers, bobcats, and

TAXONOMIC ACCOUNTS / ORDER RODENTIA 265 house cats all catch them on occasion (Pol- ple catches of red-backed voles on St. Martin lack, 1951; Voigt et al., 1976; Long and Kill- Island, Lake Michigan. ingley, 1983). Probably short-tailed shrews One specimen (UWSP 1040) had a belt- (Blarina) eat them. ed white color pattern, from Waupaca, Wis- Tapeworms, fleas, sucking lice, mites, consin. It showed the ventral white extending chiggers, and ticks infest these mice (Jack- fairly continuously forward to each manus. son, 1961; Manville, 1949; Timm, 1975; Geographic variation. Colors are vari- Scharf and Stewart 1980; Merrit 1981). Be- able, and large samples are essential to coming forest mammals they are often devastat- pare color from place to place. The variabili- ed by forest fires (Wrigley, 1975; Krefting and ty is constant geographically. The voles are Ahlgren, 1974). brighter in winter. Home Range and Density. Nothing has Additional Natural History. Merritt (1981) been done on ascertaining the home range reviewed natural history of the red-backed vole. size of this species in Wisconsin. Manville Geographic Variation. No geographic (1949) in Upper Michigan found a high den- variation was observed in the red-backed vole. sity of 4.4 per acre in white cedar habitat. He Specimens examined. Total, 296. Adams, found that females were territorial but males Ashland, Bayfield, Clark, Columbia Co.: not so much. Occasionally red-backed voles Portage, 1. Dane Co.: Town of Burke, Madi- aggregate in groups for winter. Robinson and son, 1. Door Co.: Washington Island, Ped- Werner (1975) found 6 to 8 voles per acre in ants Lane, 14. Douglas, Dunn, Florence, Upper Michigan. In Minnesota Gunderson Fond du Lac, Forest, Iron, Jackson, Juneau, (1962) found home range of about one third Langlade, Lincoln, Manitowoc, Marathon, of an acre. In winter Beer (1961) found that Marinette, Marquette, Oneida, Ozaukee, voles kept to narrow, well-packed paths in Portage, Sawyer, Shawano, Sheboygan, Tay- home ranges about 0.25 acres. They seldom lor, Trempeleau, Vilas, Washburn, Washing- left the security of a brush pile or fallen log ton, Waupaca, Waushara, Wood counties. (of 17 mice recaptured). Blair (1941) found Other records. Jackson 1961; Pitts, home ranges in the growing season as large 1983. Ashland Co.: All Apostle islands ex- as 3.6 acres for a mouse caught 22 times. cept Madeline and Long isles (Kantak, 1981). No difference was found for home range size Monroe County; Johnson (1978). Wide- of males or females. spread in Upper Michigan, see Baker 1983. Remarks. For remarks on the relation between Clethrionomys and Microtus pennsylvanicus, see account of the latter species. Genus Pitymys McMurtrie The red-backed vole is often taken with masked and short-tailed shrews (e.g., Timm, 1975). In Pitymys the lower first and upper third The red-backed vole is seldom abundant but molars are primitive. The M/1 has the ante- reaches high densities in some years. On Wash- rior island slightly constricted, with only two ington Island, since the invasion of white-foot- to three salient angles between it and the ed mice, red-backed voles seem scarce. posterior loop. The upper third molar has only Red-backed voles are active day or night, two islands (two closed salient angles) between winter or summer (Merritt, 1981), and readi- anterior and posterior loops. ly climb trees. Populations may have irregu- The close resemblance of the complex lar multi-annual cycles. Red-backed voles are molar teeth in P. ochrogaster and P. pine- thought to be solitary but Pitts (1983) caught torum persuades me to combine the two spe- six adults in the same tunnel beneath a de- cies in Pitymys. Repenning (1983) included cayed stump. Long (1976) also made multi- Microtus quasiater for the same reason. Pa-

266 THE WILD MAMMALS OF WISCONSIN pers bv Pfaff (1990) support this combination. complex cheek teeth, nor the paleontological Pfaff found both Pedomys and Pitymys dis- origin of Pedomys and Pitymys, Shenbrot and tinct back into the Irvingtonian, middle or ear- Krasnov (2005) say “molecular data” support ly Pleistocene Epoch. This age suggests they the narrow interpretation of the subgenus. may comprise a valid genus. DeCoursey (1957) Pedomys is somewhat related in molecular had difficulty separating fossil ochrogaster and characters (not to M. xanthognathus). pinetorum even as species. He relied on such characters as enamel thickness, shape of the lower m/3s, and the variable distance between Pitymys ochrogaster (Wagner) re-entrant angles (three labial and four lingual). Prairie Voles Although teeth in American Pitymys differ from those of some Microtus, their simple 1842. Hypudaeus ochrogaster Wagner. In Schreber. pattern (Carleton, 1985) might support includ- Die Saugethiere... Suppl. 3: 592, type from ing Pitymys in a catch-all arrangement of Mi- America, probably New Harmony, Indiana crotus. Zakrewski (1985) found Pitymys and (Bole and Moulthrop, Publs. Cleveland Mus. Pedomys (P. ochrogaster) congeneric. Alloz- Nat. Hist. 5: 157,1942). yme variation (Modi, 1987; Moore and Jan- 1853. Arvicola austerus Le Conte. Proc. Acad. Nat. ecek, 1990), which provides different infor- Sci., Philadelphia, 6: 405, type from Racine, mation than dentition, also shows quasiater Wisconsin. This name is a junior synonym. related to ochrogaster. Two diagrams using 1898. Microtus (Pedomys) ochrogaster: J. A. Allen. the same isozyme matrix, the Rogers’ Genetic Bull. Amer. Mus. Nat. Hist., 10: 459. Distance dendrogram and the Unrooted Tree 1966. Pitymys (Pedomys) ochragaster: Ellermann of Fitch and Margoliash, illustrate contrasting and Morrison-Scott. Checklist of Palearctic relationships. The RGD tree shows Pitymys and Indian Mammals. British Museum Nat. pinetorum and P. ochrogaster in a related Hist., P. 681. cluster if M. californicus and M. quasiater are 1990. Pitymys ochrogaster: Long. The voles and bog included. This cluster is a subgenus if Micro- lemmings of Wisconsin. Trans. Wisconsin Acad. tus contains the entire assemblage. Sci., Arts & Letters, 78: 88. See also Zakrzews- The Fitch-Margoliash diagram shows P. ki, 1985, The fossil record [of microtines]. Mono- ochrogaster and M. (= Pitymys) quasiater graphs Amer. Soc. Mammalogists, No. 8. related, but apart from a branch pinetorum and Microtus californicus. This suggests a This binomen means ochraceous-bellied basic difference between the two Pitymys mouse. Because they are found in grasslands branches, which is reflected by their habits and they are called prairie voles. Many workers morphology. I think it argues for inclusion of follow Hall (1981) and a few others in assign- both branches in a larger grouping, either in a ing these voles to the diverse genus Microtus separate genus Pitymys, or within the Micro- [When diverse genera are assigned to this ge- tus. One might also argue for four separate nus, their overlaps allow assignment of other genera from these charts. And one might ar- divergent genera to it, creating a sort of a gue for several arvicoline (microtine) genera variable catch-all taxon.] Usually workers fo- based on morphological characters. In six of cus attention on the distinctness of prairie six phylogenetic dendrograms, Modi (1987) voles by use of the subgenus Pedomys. I try found P. pinetorum and P. ochrogaster re- to call attention to the close similarity of the lated, and Microtus californicus also close to dentition of P. ochrogaster with that of the them. In one of the six, pinetorum and ochro- pine vole Pitymys pinetorum. Assigning gaster were set apart from the other voles. them to the genus Microtus does not accom- With no consideration of the congruent and plish that. The complex molar teeth both in

TAXONOMIC ACCOUNTS / ORDER RODENTIA 267 the upper and lower jaws are practically iden- inal teats are used to nurse a litter. A hip gland tical in the two species. That surely means prominent in M. pennsylvanicus is small or common ancestry, which the use of Micro- absent (Stalling, 1990). tus obscures. The color is generally a brownish gray Description. The prairie vole is stout and with a rusty tone in the dorsal fur, which is slightly grizzled, having slightly rusty ochra- flecked with blackish and buffy-ochraceous tips ceous brown intermixed dorsally with buff, of hairs. The sides are similar to the mid-dor- black, and gray hairs. The venter is usually sum but paler, and the venter is gray washed overlain with bright ochraceous. The tail is of with cinnamon-ochraceous or even orange or medium length, the ears short, and the eyes rarely rusty In young animals this ochraceous bright brown. The skull is more or less round- color, on the venter, is hardly developed, and ed in adults, but the braincase becomes more the belly seems gray, but the coarse dorsal angular as the animals age. The auditory bul- fur is distinctively gray (in M. pennsylvanicus lae are moderately inflated. The upper inci- the young are exceptionally dark brown with sors lack grooves. The vole-pattern of prism a slightly reddish tint). Hind feet are buffy development on the upper molars shows typ- brown with 5-6 tubercles on each. The bog ical anterior loops, posterior loops, and in- lemming Synaptomys also has coarse red- tervening prisms, all of which are dentine is- brown fur but its tail is remarkably short and lands surrounded by enamel walls. The mid- its upper iincisors are slightly grooved (see ac- dle upper molar lacks the distinctive minute count of that genus). There is an annual molt loop or islet posteriorly, which characterizes in summer in P. ochrogaster, but it may oc- the common Microtus pennsylvanicus. The cur (in a duration of three weeks) anytime in posterior upper molar has fewer prisms (only the year according to Stalling (1990). The molt two closed triangles, and with an elongate c- begins pectorally, spreads across the under- shaped posterior loop) than M. pennsylvani- sides and upwards, and ends on the dorsum. cus, and the lower first molar shows a minor There are two races differing significant- constriction (not entirely closed off) of the ly in size in Wisconsin. The males average anterior loop resulting in fewer prisms than slightly larger than females, but the sexes are in M. pennsylvanicus. The lower third molar similar in size according to Stalling (1990). has three transverse loops with continuous See accounts for each subspecies below. dentine (i.e., no closed triangles). The teeth, Dental formula. I 1/1, C 0/0, P 0/0, M regardless of complexity, closely resemble 3/3 = 16. those of Pitymys pinetorum. A skull figure Geographic Distribution. Large prairie precedes Muridae. voles have ranged northward from the Illinois The cranium of P. ochrogaster is much prairies into the southern and southwestern narrower than that of P. pinetorum. See ac- grasslands of Wisconsin, usually dwelling on count of the pine (or woodland) vole for addi- the dry, well-drained slopes of the southern tional characters. The baculum of P. ochro- uplands. The minor vole ranges along the gaster has a basal stalk and three minute glacial sands left by the Wisconsin glacier, and prongs extending into the glans penis. It mea- occurs in low numbers in a band from Stevens sures 3.2-4 mm in length (Stalling,1990). The Point, across the Wisconsin River westward, chromosomes number 2N = 54, FN = 64 as far south as the Necedah National Refuge, (Hsu and Benirschke, 1971). and far westward of Wisconsin, always show- There are six mammae, but some have ing small size and dwelling in bleak, sandy reported the typical Microtus eight. In close- habitat. Possibly this kind is one of the few ly related P. pinetorum there are only four. Wisconsin taxa with western origin, which Often in P. ochrogaster only the four abdom- immigrated eastward into Wisconsin, or inva-

268 THE WILD MAMMALS OF WISCONSIN sions may have ranged northward along re- no longer there. This race (or perhaps a spe- ceding glaciers in several states. cies) seems destined to vanish from Wiscon- Status. This species is fairly abundant in sin before we know about it. some of the southern counties where the soil Habitat. The prairie vole prefers dry prai- is sandy loam, usually on the highlands with rie or old fields with more or less sparse veg- sparser vegetation and in scattered cedars and etation including Andropogon, Panicum, prickly pear. It often comes into contact with Sorgastrum, Poa, forbs, lichens, some brush, Microtus pennsylvanicus although the latter such as wild plum, hazelnut, brambles, jack prefers more moisture in its diet and habitat. pine, and other short, stubby grasses. Occa- The prairie vole is important in the food chains sionally the dry habitats merge with more lush of carnivores, raptors, snakes, and other pred- vegetation, dense grassland, alfalfa, and so ators that live in the high, dry prairies. It feeds forth. Microtus pennsylvanicus may co-ex- on weed seeds and on some insects. Prairie ist with prairie voles in such dry-wet grass- voles do not make much impact on agricul- lands. In general the prairie voles inhabit west- ture in Wisconsin according to Martin (1956), ern and southern grasslands in Wisconsin, Crawford (1971) and Jackson (1961). following along unplowed railroad rights-of- The larger prairie voles in the southern way, roadsides, and ranging into jack pine- subspecies (P. o. ochrogaster) are not in any oak savannas, cedar glades, and weedy fields. apparent peril, although herbicides and de- This vole often makes its surface runways struction of the prairies themselves may dec- and burrow systems in soils of sandy blow- imate and eradicate some future populations. outs and sandy prairies, on well-drained high- As for the minor prairie vole, which does not lands and weedy slopes, and in cut-over grain intergrade with the other prairie vole in Wis- fields, hazelnut groves, and occasionally open consin, and which may be a valid species, it is meadows and oak groves. It avoids wet plac- perfectly harmless and is in great peril. In cen- es such as marshes, seldom occurs in woods, tral Wisconsin it seems to have been eradi- and never in forest. Jackson (1961) stated that cated, not so much by agriculture but by de- the preferred habitat was “native prairie sod.” liberately combining plowing and fire to con- Extending all about the home are numer- trol vegetation and prevent wild fires along ous surface runways, which are conspicuous railroad tracks. The known populations are paths, where obstacles such as large stones are avoided, plants cut off, and the trails are occasionally rutted or trenched in the soil. Piles of cuttings are often evident. A runway may extend as far as 70-80 feet (22-24 m), and occasionally farther. They average about two inches (5 cm) in width. Many show signs of excavation as ruts. One or several openings lead into underground tunnels (Getz and Cart- er, 1996). Grass cuttings are often used for the nests and found with food caches (Hoffmeister, 1989). Getz and Carter (1996) describe the nest as approximately 10 to 15 cm in diameter, comprised of tightly interwo- ven and finely shredded grass. Materials are opportunistically used to protect nests, such Photo of central Wisconsin Pitymys ochrogaster minor. as a piece of plywood left in a field, bark or This rare mammal’s population was eradicated. rocks. Nests occasionally are found above

TAXONOMIC ACCOUNTS / ORDER RODENTIA 269 ground. Prairie voles pile excavated dirt over a nest site forming a mound (Getz and Hof- mann, 1986). The tunnel entrance to the underground nest is about one and one half inches (3.6 cm) in diameter. Getz and Carter (1996) recorded a diameter of approximately 3 cm. Often excavated soil is thrown out the entrance. The tunnel drops nearly straight down, and is more easily dug in sandy soil. Its depth ranges from “a dozen or more centimeters” below the surface. Jackson (1961: 239) described a nest in Sauk County made of barley

Maps showing geographic distribution of Pitymys (= Microtus) ochrogaster in Wisconsin and North America.

270 THE WILD MAMMALS OF WISCONSIN grasses, 5 1/2 inches (138 mm) in greatest In Illinois the prairie vole has young in diameter by four inches wide by three inches any month, but none was observed pregnant depth. The entrance into the nest chamber in August or January by Hoffmeister (1989). was about 7/8 inches (24 mm) in diameter. In 330 adult females, 94 contained embryos Foods. Cole and Batzli (1979) found that (mean 3.67, 1-8) according to Hoffmeister. succulent forbs were required in the diet of the Cole (1977) found 56 females in bluegrass prairie vole. Cole (1977) had previously report- habitat to average 3.66 embryos; in alfalfa ed that prairie voles eat clovers (Trifolium sp.), the average litter size was 5.03 (N = 30) in dandelions (Taraxacum), alfalfa, Penstemon, summer and in autumn 5.19 (N = 21). Ambrosia, Solidago, Poa pratensis, some tim- A chemical signal, a pheromone, in the othy (Phleum), and big bluestem (Andro- male’s urine stimulates estrus in virgin females. pogon) in Illinois. In winter, Long (1968b) The pheromone is usually received from an found wild plum seeds cached and eaten, in unfamiliar (probably not closely related) male central Illinois. Each seed was opened the same by sniffing of the genital area. In two days she way on the same side. Roots and mosses were achieves estrus and mates. The female usually eaten in winter and spring, and Long also re- has a post-partum estrus, mating again and ported gnawed roots. Fisher (1945) found large usually with her original mate (Getz and Cart- caches of horsenettles, and Criddle (1926) er, 1996; Carter and Getz, 1993; Hofmann found caches of wheat, rye, and oats made by et al., 1984; and McGuire and Getz, 1991). P. o. minor. In Indiana important foods were Growth was discussed for captive prairie bluegrass, red clover, lespedeza, tumblegrass, voles by Hoffmeister and Getz (1968). Skulls and roots. Energy budgets were studied by Fle- are judged adult when the basioccipital-ba- harty and Mares (1973). sisphenoid suture closes. In old adults the cra- Getz and Carter (1996) suggest that es- nium becomes angular in dorsal view. sential forbs are not always available to prai- Mortality. Prairie fires kill many prairie rie voles. Famine affects the social system (see voles in Wisconsin, and may destroy the hab- below) and cyclic densities. itat in thin, sandy soils with little vegetation, Reproduction. In southern Wisconsin the preventing re-establishment of the vole pop- praire vole may breed most of the year, but ulations. Hard winters might destroy some little if any in winter. A female may produce populations of voles in Wisconsin, for the prai- several litters in a year. Gestation is about 20- ries suffer deep frost where the winds clear 21 days. Litter size is about three or four (range off the snow. Predators that feed on prairie 2-6). Young weigh 2-3 g at birth, and are born voles include several kinds of raptors of prai- naked with eyes closed. Pelage grows rapidly rie and savanna (Stalling, 1990). These in- and the eyes open after 9-10 days. By the 17th clude kestrels, marsh hawks, red-tailed and day, the young are furred and can forage for rough-legged hawks, and prairie dwelling spe- food (Fitch, 1957). The young can breed in cies of owls. Weasels, striped skunks, badgers, 36-40 days perhaps even in 26 days (Stalling, foxes, and coyotes eat them, and rattlesnakes, 1990). According to him, the young are born fox snakes and bull snakes catch a few. Gar- 30-35 mm in length, with eyes closed. The ter snakes probably eat young voles. only records of reproduction I have from Wis- Getz et al. (1990) discussed the impor- consin are as follows: P. o. minor lactation 22 tant impact of snakes on numbers and social July, 2 subadults Sept. 25 and 20 October; P. organization of prairie voles in Illinois. In Wis- o. ochrogaster testis 12 mm on 3 July, young consin snakes are not abundant, not in the on 1 July and 1 November. The voles usually central parts of Wisconsin certainly, and they live less than a year, but Getz (1965) kept one probably enter hibernation sooner. The com- in captivity for 35 months. petition with Microtus pennsylvanicus often

TAXONOMIC ACCOUNTS / ORDER RODENTIA 271 drives the prairie voles from lush vegetation Remarks. Active day or night, especially onto bleak sandy soils where runways are less before dawn or darkness, the prairie vole goes protective (bordered with thin growths of grass- about its activities of feeding, nesting, and es). The voles may have less contact with forbs, breeding. It is often (exceeding half the breed- berries, and other foods. Drought and loss of ing adults) monogamous, and stays mated for available water also play a role in mortality. relatively long periods (Getz and Hofmann Parasites include five species of fleas, one 1986). These sociable and “comical” mice chewing louse, six mites, and internal trematodes, threaten, box, wrestle, and play, but with lit- cestodes, and nematodes (Jackson, 1961). Timm tle actual fighting. They are not so likely to (1985) listed cestodes, spiny-headed worms, kill one another, or to foul their surround- mites, ticks, lice, and fleas. Little is know about ings, as are captive M. pennsylvanicus. Hav- diseases in these voles, but the numerous ecto- ing less need for water (Getz, 1963) the ani- parasites possibly transmit some diseases to hu- mals seem cleaner and in my care made ex- mans and other mammals on contact. cellent pets. Jackson (1961) also believed prai- Home Range and Density. The home rie voles in Wisconsin were more colonial and range (Jackson, 1961) is between 0.1 and 0.2 sociable than meadow voles. In my experi- acres. Swilhart and Slade (1989) found males ence that is true, in either P. o. ochrogaster to utilize much larger home ranges than fe- or diminutive P.o. minor of central Wiscon- males which suggests a promiscuous mating sin. Although they threaten one another by system. Stalling (1990) gives home range val- standing as tall as possible and gaping, there ues as 0.11-0.22/ha. Usually the home range, is little biting. Often they cuddle together. larger in males, is less than 0.1 ha. Density Thomas and Birney (1979) described pa- may be as high as 20 per acre. rental care and the mating system in this spe- Getz et al. (1979) found peak densities cies for Minnesota. The social behavior and of the larger prairie voles in Illinois in alfalfa mating system have been well studied in central mixed with goldenrod and grasses, as many Illinois by Getz (1993), Carter and Getz (1993), as 240 mice per ha in November. Lowell Getz Getz et al. (1992), Getz et al. (1993), Hofmann (personal correspondence) told me that the et al. (1984), McGuire et al. (1993); McGuire et number was as high as 638 per ha in 1985. al. (1990), and Getz and Carter (1996). They found only 110 in July 1976. In 1974, Briefly the mating and social behavior is there were fewer than 10 per ha. In some sketched as follows. Based on 25 years of lab- populations there are only annual cycles (Getz oratory and field studies and monitoring of et al. 1987). The presence of some other 850 free-living social groups of prairie voles, species seems to decrease prairie vole abun- Lowell Getz and his associates found several dance, as Sigmodon in Kansas, and M. penn- mating systems that showed variance due to sylvanicus in Illinois (Stalling, 1990). Where certain factors. These were chiefly the quan- vegetation is sparse the voles are fewer (Fitch, tity of foods and the population density. In a 1958, in Kansas; Mumford 1969, in Indiana). species known for periods of low and remark- Getz et al. (1981) found prairie voles dis- ably high abundance, sometimes cyclic (i.e., persing along railroad rights-of-way. The pop- multi-annual), an abundant food supply per- ulations of P. o. minor that formerly occurred mits population growth, whereas mortality, in Portage County, Wisconsin, were either in especially predation, lowers density. such areas or near them. The minor subspe- The main social group is the monoga- cies seems never to have been common in mous pair, with or without young. The paired central Wisconsin. It was local in distribution, adults stay together, sharing nest and home and was also found on thin sandy soils or in range. The male contributes “considerable time sand barrens near dry prairie near Necedah. and effort” constructing and maintaining a nest,

272 THE WILD MAMMALS OF WISCONSIN helping groom the young, and retrieving them philopatry are adaptive in keeping young voles when they wander from the nest. Unfamiliar secure at home where food and mates are adult males are excluded from the nest. Three- easier to find in times of plenty. When food is quarters of such pairs persist until one mate abundant, monogamy is not strictly practiced. dies. A widow remains alone, or stays with Geographic variation. The two distinct young. Thus, the single female consitutes a races do not overlap or intergrade; they are second, derived social group. A widower wan- allopatric in Wisconsin. Habitat differences ders away from the nest. Between 24-45 per- and intervening woodlands and river bottoms cent of the small adult voles wander unattached. keep the two kinds apart. Accounts of the The third group is communal, an extremely two subspecies are below. Information above interesting “extended family”, comprised is for the species. mainly of the breeding adults and their offspring. These may have grown to maturity, remain in the natal nest, and may help in the Pitymys ochrogaster ochrogaster (Wagner) care of the young. Such a group is “philopat- Prairie Vole ric.” Getz and Carter (1996) report that 68 percent of males and 73 percent of females 1842. Hypudaeus ochrogaster Wagner. In remain “at their natal nests.” Males and fe- Schreber’s Saugthiere... Suppl. 3: 592.Type males that seem “unrelated” may join the ex- America, possibly New Harmony Indiana. tended family. However, 70 percent of the 1898. Microtus (Pedomys) ochrogaster: J. A. Allen. adults, in addition to the parents, are philopat- Bull Amer. Mus. Nat. Hist., 10: 549. ric offspring (that stayed home). Of wander- 1853. Arvicola austerus Le Conte. Proc. Acad. Nat. ers, the unattached female usually finds wel- Sci. Philadelphia, 6: 405. Type from Racine, come in a nest, either as a single or joining in Wisconsin. a pair. Group members seldom (not more than 4 percent) switch to other groups. Description. See account of the species. The social groups vary in size seasonal- For comparison with P. o. minor, see account ly. Communal living increases in mid-Octo- of that race. Some measurements in mm of ber (in central Illinois) and remains high in three specimens from Lynxville average as winter. The group size averages 8 voles at follows: Total length, 153 (152-155); length these times, and often may exceed a dozen. of tail, 39 (38-40); hind foot length 18.7 (18- After suffering winter mortality, the survivors 19); greatest length of the skull 26.8 (25.8- of various communal groups pair anew, mix- 26.9); length of the maxillary tooth-row 6 ing members of different families. Incest in (5.6-6.5); zygomatic breadth 15 (14.9-15.1); families of monogamous parents is prevent- interorbital breadth 4 (3.95-4.1); cranial depth ed, and in communal groups there is mostly 9.6 (9.6-9.7); and length of nasals 7.55 (7.5- outbreeding. This rules out inbreeding, which 7.6), and weights 47 (44-50) g. is often a cause of a communal clan system, Geographic Distribution and Status. See with breeding restricted to one or two pairs. account of the species. See Map. Another cause for such a system is patchy Specimens examined. Total, 23. Craw- environment. Both the runway-system, ex- ford Co.: Lynxville, 3 USNM. Dane Co.: tending in branching patterns that do not fill Beeny, Sec. 18, T8N, R7E, 1 UW-Madison. 5 the areal space, and the lack of sufficient foods mi. N Cross Plains, 1 UW-Madison. Dodge to feed large populations could account for Co.: Beaver Dam, 1. Rock Co.: Milton 5 UW- such an extended family system. Getz and Madison. Sauk Co.: Pleasant Valley Road, near Carter (1996) discuss the latter, i.e., when Sumpter Church, 3. 1/4-3/4 mi,. S, 2 mi. W forbs may be in short supply. Monogamy and Prairie de Sac, 3. 4 mi. W Prairie de Sac 4.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 273 Pitymys ochrogaster minor (Merriam) persists on thin, sandy soils. When plowed Minor Prairie Vole into the sand, the habitat is destroyed. Habitat. Wydeven and Wydeven (un- 1888. Arvicola austerus minor Merriam. Amer. Nat. published report, 1976) collected P. ochro- 22: 6. Type from Turtle Mountains, lower gaster minor and M. pennsylvanicus to- slopes, Bottineau Co., North Dakota. gether at a sandy habitat near the Little Plo- 1942. Microtus ochrogaster minor: Bole and ver River, called the Stream Lab. The mead- Moulthrop. Science Publs. Cleveland Museum ow vole preferred the lush and dense vege- Nat. Hist., 5: 160. tation. After the grasses were burned off 1990. Pitymys ochrogaster minor: Long. Trans. the land was plowed with the result that Wisconsin Acad. Sci., Arts and Letters, 78: 94. these voles vanished. Eradication also was the case along the Soo Railroad tracks at Description. There is only one signifi- the edge of Stevens Point, near the Coun- cant difference noted between the two voles, try Club, and the property later was devel- that of size. Their behavior and habitat pref- oped (sodded, fenced). After fire and plow- erence are also somewhat different. The ing this population also vanished from the auditory bullae are relatively more inflated right of way. in P. o. minor, although the opposite has Specimens examined. Total, 29. Clark been reported. Co.: Brick Creek, near Owen-Withee, 1. Fos- Not one large vole has been found in center Twnsp., 2 UW-Madison. No specific lo- tral Wisconsin. That is highly significant. The cality 1. Juneau Co.: 4.5 mi. N, 1 mi. W cranial measurements in mm of six adults from Necedah, 2. Portage Co.: Stevens Point, 14. Stevens Point are 24.6 (24.1-25.6) for great- Whiting, 2. Plover, 5. 8 mi. S Plover, 1. est length of the skull, zygomatic breadth 13.7 Waushara Co.: Saxeville, 1 UW-Madison. (13.5-13.9), interorbital breadth 3.95 (3.8- 4.0), and length of the nasals 6.59 (6.7-7.3). One specimen from Clark County was, re- Pitymys pinetorum (Le Conte) spectively, 25.9, 13.4, 3.8, and 7.0 mm. Pine Vole or Woodland Vole Geographic Distribution. See Map. Cory (1912) speculated on this little vole’s presence 1830. Psammomys pinetorum Le Conte. Ann. Lyc. in Wisconsin. It escaped notice until I began Nat. Hist., New York . 3: 133. Type from pine catching them, with never an adult obtained forests of Georgia, probably near the Le Con- as large as would be found in the southern sub- te Plantation near Riceboro. species (see Long, 1976). The race seems to 1841. Arvicola scalopsoides Audubon and Bach- have extended its range on glacial sands from man. Viviparous Quadrupeds of North Ameri- Minnesota, where it also is imperiled, espe- ca. p. 97. Type from Long Island, New York. cially by land use. Intergradation between the 1896. Microtus pinetorum: Miller. N. Amer. Fau- large and small voles was once reported, since na, 12: 9. disputed, in southeast Minnesota, which in- 1858. Arvicola kennicotti Baird. Mammals. In Re- tergradation ought to be studied. There is no ports Explor. Survey... 8(l): 547. Type from sign of it in Wisconsin. The two races approach Illinois (see Long, 1990, for discussion of spec- one another in two areas, but are separated imens examined by Baird). by about 40 miles (64 km). 1894. Arvicola pinetorum: Ellermann and Butler. Status. This little vole is in grave peril. Proc. Indiana Acad. Sci., p. 127. No specimens have been taken in central 1966. Pitymys pinetorum: Ellermann and Morrison- Wisconsin in over ten years. Plowing the hab- Scott. Checklist of Palearctic and Indian Mam- itat is extremely detrimental to this vole, which mals. British Museum (Nat. Hist.), London.

274 THE WILD MAMMALS OF WISCONSIN 1990. Pitymys pinetorum: Long. Trans. Wisconsin woodlands, respectively. The pine vole also Acad. Sci., Arts and Letters. superficially resembles the red-backed vole (see account of that species) and the bog lemming, The binomen means a mouse from pine which has coarser, grizzled fur and a groove forests, but seldom is this mouse found in them. on each upper incisor (see account of that spe- For that reason in recent years the name cies). The western race, P. p. nemoralis is red- “woodland vole” is often used for a common dish and comparatively large, judging not only name. The name of the synonym P. p. schmid- from the single specimen Jackson reported, ti honors an amateur naturalist, F. J. W. but based on numerous specimens examined Schmidt, from Clark County, Wisconsin, and from the race itself. The pine voles in central the name kennicotti honors a naturalist, Rob- and southern Wisconsin (P. p. kennicotti) are ert Kennicott, from the Chicago region. duller in color, even in pelage showing its max- Description. The body is modified for imum coloration between molts. The dull fossorial life, in that the head is broad, the brownish voles do show a reddish tint, more forefeet and foreclaws well developed, the tail chestnut than red, but a dull walnut (purplish short for turning about in the burrow, and as in some walnut wood) brown is the domi- the fur is soft and fleecy, said to be “mole nant tint. That color becomes constant south- like.” The behavior is likewise curious and ward into Illinois. Baird examined, and his fossorial, for the mice seldom leave the tun- name was based upon, some of the same pine nels, throw up dirt in the fashion of moles, voles that I examined on loan from the Chica- and are exceptionally difficult to catch. (I have go Natural History Society collection. They tried and never caught one.) Although the differ from the reddish voles westward and teeth are similar to those of P. ochrogaster eastward. The name by Baird seems appropri- (see account of that species), the skull is so ate; the name by Jackson is a junior synonym. much broader it can be identified (see key to I have no idea what color the pine or wood- voles and bog lemmings in Microtinae). land voles are in southern Michigan. Reportedly there are six tubercles on the A darker fur is replaced in May or June by hind foot. Chromosomes number 2N = 62, FN a lighter fur, and a winter molt begins in No- = 62 (Hoffmeister, 1989). There is a report of vember (Smolen, 1981). According to Reich eight mammae; most agree to six. The bacu- (1981) the annual molt is mostly of underfur. lum’s stalk has three distal calcified projections. For P. p. scalopsoides from eastern The dentition is discussed under the ac- states, some measurements in millimeters are count of P. ochrogaster, and in the key. Al- as follows: Total length, 121 (110-133), tail though the teeth are similar in P. ochrogaster length (very short) 21.3 (16-24.5), hind foot and P. pinetorum, they occur in different hab- 16.6 (14.5-18) where N is 25. For Wisconsin itats, i.e., prairie grasslands and deciduous voles some external measurements are from Clark County (N = 4), as follows: 119-127 in total length, length of tail 20-23, and in length of the hind foot, 16.5-18. They are 110-126, 14-16, and 16-18 from Wood Co. (N = 3), and one from Sauk Co. 109, 17, and 15, respectively. Some cranial measurements for three specimens from Wood County are in mm as follows: greatest length of the skull mean 25.1 (range 23.1-26.2), zygomatic breadth 14.1 (range 13.7-16.0), and interor- Photo of Pitymys pinetorum. By Karl H. Maslowski. bital breadth 4.6 mean (range 4.5-4.75).

TAXONOMIC ACCOUNTS / ORDER RODENTIA 275 Dental formula. I 1/1, C 0/0, P 0/0, M 3/3 = 16. Geographic Distribution. Found locally in leaf litter, with some clay in the soil, and probably with affinity to relict hickory-maple woodlands. Pine voles seem confined to hardwoods or mixed hardwoods in the southern half of the state. Occasionally they range from the thick humus and clay-loam soils of the woodlands into weedy pastures, road rights-of-way, or orchards. A western race has managed to cross the Mis- sissippi River into western Wisconsin, but so far

Maps showing distribution of Pitymys pinetorum in Wisconsin and North America.

276 THE WILD MAMMALS OF WISCONSIN is known from a single specimen. The more seen, and from beech and hemlock woods, also widespread race has extended its range north- which I have not seen. He also mentions sandy ward in the past as far as Clark and Wood coun- soils and brushland (see also Pool, 1932; Jame- ties, and long ago as far as the Door Peninsula. son, 1949). The tunnels and mound systems I Today it is found in very few localities. There is have observed require clay to resist collapse. a questionable record from Brown County, The pine vole occurs in a wide range of habi- which Jackson (1961) reported as pinetorum tats outside Wisconsin (Paul 1970). It is most (based on a lost specimen in the Neville Muse- commonly taken in hardwoods (maple, hicko- um, in Green Bay). See Map. ry, oak) forests but in southwest Wisconsin Status. As mentioned in the Distribution, numerous burrows were found on a dry, grassy the populations are local and small. Possibly hillside. In Clark and Wood counties the bur- all populations are relicts from an earlier warm rows ran under maple leaf litter in gray sandy- Holocene time period. The vole is beautiful, clay loam soils that roofed the burrows. When and in Wisconsin seems to live apart from abundant, meadow voles may replace Pitymys our agriculture. It follows that the Department in competition (Paul 1970). of Natural Resources Bureau of Endangered This burrowing rodent approaches in Resources might want to takes steps to pre- behavior that of the moles (see Talpidae) as it serve this endangered species. In a survey of excavates tunnels and throws up hills in fria- nuisance voles, of those that do humankind ble soil under thick leafy litter. Usually the the most damage, the pine vole was rated the burrow extends immediately below the sur- worst in the United States, especially injuri- face, raising up the soil above it in a mole-like ous in orchards. It is a difficult problem to run. It may descend as low as several inches justify preserving such a pest. In Wisconsin [~75 mm], and occasionally as deep as a foot the pine vole is so interesting a mammal, fill- [0.3 m]. The nest cavity may be an enlarge- ing a niche like some sort of forest-dwelling ment of the tunnel or in a side branch often mole, and so uncommon, it might well be protected above by a stump or tree root. Dry preserved. No species is bad everywhere. In leaves and grasses are wadded into the tun- Wisconsin, the species probably fills a useful nel, and two or three entrances penetrate this role in the food chains of deciduous wood- nest. Schmidt (1931) describes the nests of land predators. It makes soil, kills insects, and voles from Clark County as follows: eats plant parts when in that kind of habitat. “Two nests were located... two inches below the Habitat. Pine [= Woodland] voles prefer surface... traps set at the entrances of the nests caught deciduous woodlands with clay soils (not much both adults and half grown young. These nests had sand), covered with thick leaf litter and mold, an outside measurement of 100 mm, and were con- under which the dirt from tunnels and hills structed of maple leaves... The surface burrows of pine aand excavations are evident. The fallen leaves voles were constructed just below the leaves, and they are primarily maple with hickory observed at often form a network covering several square yards. all localities I have seen in Wisconsin (Schmidt, From these burrows under the leaves other burrows 1931; Long 1974; Long, 1990). These voles penetrate the soil. Dirt brought up from below is left also range into overgrown fields not far from under the leaves. This distinguishes the mounds from their woodland sources. The distribution, as those of Condylura cristata, which are piled up on mentioned above, is local and many suitable top of the leaves or grass... In diameter the burrows habitats seem uninhabited by these voles. of pine mice are 30-35 mm wide and 25-30 mm high. They are difficult to catch in traps. Pine mice burrow both day and night... Mice caught Jackson (1961) reports pine voles from shoving out dirt were always coming out head first.” grazed red and white oak woodlots having fri- Foods. Smolen (1990) lists foods of pine able top soil, a habitat I have personally not voles in New York and Pennsylvania as grass

TAXONOMIC ACCOUNTS / ORDER RODENTIA 277 roots and stems in summer, fruit and seeds in days young are active, well furred, and the eyes autumn, and bark and roots in winter. Denti- are beginning to open. Eyes open by 12 days, tion of the pine vole not only resembles that of and the young are weaned from 16-24 days. P. ochrogaster, but the foods do likewise, even The young weighs then about 15 g, almost 2/ from different habitats. Forbs and shoots of 3 the adult body weight. Males begin breeding grasses probably were the foods eaten in spring. at about 7 weeks, and females in 10-11 weeks. No doubt these voles also feed on grubs, and There seems to be a postpartum estrus. perhaps worms. Worms are difficult to find in The young pine voles can be aged by the stomach contents, but seem likely for a mole- weights of the eye lens (Gourley and Jannett, like mammal to eat. There may be more di- 1975). Additional information on growth and verse fare than so far discovered, but the pri- development is summarized by Smolen (1990). mary foods seem to be roots and parts of plants. Mortality. Jackson (1961) and Schmidt In the eastern states this vole emerges from its (1931) suggested carnivores such as weasels burrows to feed on bark, and causes extensive and skunks, snakes, and especially short-tailed damage to orchard trees. Schmidt (1931) re- shrews were predators of pine voles. No stud- ports tubers of Dutchman’s breeches, and an ies have been made in Wisconsin. Land use, unknown tuber as foods. Hoffmeister (1989) such as lumbering or developing a ski run in reported food contents of 13 pine voles taken their habitat at Powers Bluff, Wood County, in autumn in Indiana consisting of roots, green could be an adverse problem for that local vegetation, and mast, in 63 percent of the food population. Some known predators from else- volume (and at a frequency of 96 percent). There where include red fox, gray fox, raccoon, also was a little insect material and the fungus house cat opossum, least weasel, and mink. Endogone. When these voles are found in Smolen (1990) recently listed avian predators, grassy vegetation one presumes they eat parts barn, long-eared, short-eared, barred, screech (probably roots and bulbs) of grasses and forbs and great-horned owls, as well as red-tailed, (see Benton, 1955), and berries. Domestic broad-winged, and marsh hawks. strawberries and blackberries were eaten. Parasites include a cestode (Catenotae- Reproduction. Not much information is nia), the whipworm nematode Trichurus, and available on reproduction in Wisconsin pine a spiny-headed worm (Benton, 1955; Doran, voles, so we must rely on other states for such 1954). Fleas, sucking lice, and mites infest studies. Pine voles produce fewer and smaller the fur (see Baker, 1983). A complete listing litters than do M. pennsylvanicus according to of parasites for this species is provided by Schadler and Butterstein (1979). Apparently in Smolen (1990). Timm (1985) also reviewed Michigan the pine vole can reproduce at a high parasites including ticks, the parasitic beetle rate (Linduska, 1950). At milder latitudes the Leptinus, and the botfly Cuterebra. pine vole can sometimes breed year round (Paul, Home Range and Density. Burt (1948) 1970). Peak reproduction takes place in July reported the home range to be only one-fourth and August (Kirkpatrick and Valentine 1970). acre, and the maximal home range for an adult Hoffmeister (1989) reported Illinois pine voles male was 93 yards in greatest diameter. Actual- to breed in March and April and into Novem- ly the home range is confined for the most part ber. In Wisconsin the breeding season is proba- to the burrow system. The mice can live at least bly similar, perhaps a little shorter than this. a year (Burt 1940; Miller and Getz, 1969). Mill- Females are aggressive in courtship according er and Getz found only 19 percent of the mice to Benton (1955). Gestation is about 20-24 marked in their study in recaptures up to two days. Litter size averages about 3 (2-5 embry- months later. Being non-territorial and inhabit- os, rarely more but up to 7). At birth each young ing burrow systems, adults of both sexes and weighs about 2 g and is naked and blind. In 9 subadults mingle freely (Smolen 1990).

278 THE WILD MAMMALS OF WISCONSIN There are fluctuations in population size of Geographic Distribution. See map. Found the pine vole (Benton 1955; Gottschang 1965; in southern Wisconsin in suitable habitats. Burt 1940). Miller and Getz (1969) working in Status. See account of the species above. Connecticut found the species to fluctuate from Specimens examined. Total, 12. Clark 0 to 5.8 voles per acre over a five-year period. County. Worden Twsp. 4 USNM; 1 UWM Gettle (1975) working in Pennsylvania found (I also saw one collected by Schmidt in the voles to attain a peak of 49.2 per acre. Hamil- Leningrad (Petersburg) Zoological Institute. ton (1938) found them thriving in New York Dane Co.: Town of Vermont 1 UWM. West- orchards at 300 per acre. In Wisconsin there is port 1 UWM. Sauk Co.: Klondike Pond 2. no evidence for high densities such as these. Wood Co.: Powers Bluff 3. Remarks. In their burrow systems, pine voles Other records: Illinois: Long, 1990, in are active day or night. Schmidt (1931) found a northeast Illinois 20 specimems. young Wisconsin pine vole completely docile, but an adult he kept leaped at him and threatened to bite his finger. A curious anomaly that would Pitymys pinetorum nemoralis V. Bailey probably confuse a worker trying to identify P. Great Plains Woodland Vole pinetorum was observed by Fish and Whitaker (1971), which was a specimen having grooved 1898. Microtus pinetorum nemoralis V. Bailey. upper incisors as seen in Synaptomys cooperi. Proc. Biol. Soc. Washington, 12:89, type from Additional Natural History. Smolen (1981) Stilwell, Oklahoma. wrote a thorough review for the pine vole. 1990. Pitymys pinetorum nemoralis: Long. Trans. Geographic variation. As mentioned Wisconsin Acad. Sci., Arts and Letters, 78:90. above, there are two races, but the western one is known from a single specimen in west- Jackson (1961) reported one specimen ern Wisconsin. The races are distinctive in from Lynxville, Crawford Co., Wisconsin, both size and color. collected July 31, 1930, under leaves and dry grass close to an old brush heap on a dry hillside 50 feet above water level of the Pitymys pinetorum kennicotti Baird Mississippi River. There were many burrows Kennicott’s Pine or Woodland Vole and runways under the leaves. Two workers (Van der Meulen, 1978; Repenning, 1858. Arvicola kennicotti Baird. Mammals of the 1983) consider this large red vole to be a Explor. Pacific Railway route... Part 1 Mammals, distinct species, on criteria of size and mor- p. 547. Type from Illinois, and the specimens I phology of the teeth. Study is necessary to examined were from northeast Illinois [Chicago confirm this opinion. Nat. Hist. Soc.]. They are inseparable from the Description. The size of this pine vole synonym Microtus pinetorum schmidti Jack- is large, with massive teeth. Size and chest- son, but distinctive from P. p. scalopsoides nut-reddish brown color are the only distin- eastward and P. p. nemoralis westward. guishing features. The holotype of P. p. nem- 1912. Microtus pinetorum scalopsoides (Audubon and oralis is dark rufescent, and the topotypes Bachman). In Cory. Mammals of Wisconsin and are not only large but quite broad across the Illinois, Chicago Field Mus., 11: 222, and others. zygomata. Specimens from Minnesota and 1941. Pitymys pinetorum schmidti Jackson. Proc. Iowa have been assigned to P. pinetorum Biol. Soc. Washington, 54: 201. Type from nemoralis, and in size and color the distinc- Worden Twsp., Clark Co., Wisconsin. tive Lynxville specimen agrees closely with 1990. Pitymys pinetorum kennicotti: Long. Trans. the type and topotypes. Wisconsin Acad. Sci., Arts and Letters. 78:90.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 279 Specimen examined. Total 1. Crawford The binomen means a tiny-eared animal Co.: Lynxville, l USNM. from Pennsylvania. Description. The largest, darkest vole in Wisconsin, the meadow vole often exceeds 160 mm in length. It has a long, blackish tail, blackish gray-brown feet, and its belly is overlain with whitish, pale buff (and surprisingly, occasionally in rust, reddish, cinnamon buff, or ochraceous). The skull, rostrum, and nasals are long and narrow, the braincase well extended posteriorly. The upper third molar consists of an anterior loop, three closed prisms, and a distinct posterior crescentic Meadow Voles. Ernest Thompson Seton. loop. In the middle upper molar an extra small posterior loop is squeezed in, rarely absent in adults. The lower first molar is Genus Microtus Schrank pinched (constricted) anteriorly, so that there Meadow Vole and Allies are five closed prisms behind the anterior loop. This species is identified with certainty by the Teeth elaborate, lower first molar with deep minute loop of the middle upper molar, and constrictions forming separate islands or the identification confirmed by the extra prisms, five salient angles between anterior is- prisms in upper third and lower first molars. land and posterior loop, upper third molar elab- The skull varies to 28.6 mm in length, orate with three closed prisms (three closed the yellowish incisors projecting forward. The triangles on either side) between anterior and incisive foramina exceed 5 mm in length. The posterior crescentic loops. Microtus pennsyl- baculum is a stalk with three calcified pro- vanicus is a large, long-tailed, dark brown vole. jections. There is a slight reddish (not ochraceous) cast in the dark walnut brown underparts (especially in late summer and fall), Microtus pennsylvanicus (Ord) which are evenly colored and hardly grizzled at all. The dorsal pelage, long and lax in win- Microtus pennsylvanicus ter, is remarkably constant in color through- pennsylvanicus (Ord) out the state. Meadow Vole

1815. Mus pennsylvanica Ord. In Guthrie, A new geography 2nd Amer. Ed., vol. 2, p. 292. Type from “Meadows near Philadelphia,” Pennsylvania. 1895. M[icrotus]. Pennsylvanicus: Rhoads. Amer. Nat., 29: 940. 1841. Arvicola fulva Audubon and Bachman. Proc. Acad. Nat. Sci. Philadelphia, 1: 96. Type from a “western state”, probably Illinois. 1858. Arvicola riparia var. longipilis Baird. Reports of an Explor. Survey... Part I, Mammals, Type from West Northfield, Illinois, or Racine, Wis- consin. Skull of Microtus pennsylvanicus..

280 THE WILD MAMMALS OF WISCONSIN In ventral coloration the range of whitish The left hind foot was normal, the other three to buff to buff-ochraceous and rarely even dark- feet white. The tail was normal. The venter er rust varies remarkably, based on a scale of was pure white without gray at the hair bases. pale to dark ochraceous, one to five. The stan- There are six tubercles on the hind foot. dards are (1) specimen UWSP No. 1610, pure Chromosomes number 2N = 46, FN = 50. silvery white; (2) buffy white No. 377; (3) whit- There are eight mammae. According to Reich ish buff No. 1089; (4) buff No. 947; (5) buffy (1981) the annual molt is mostly of underfur. ochraceous No. 3190 or 2259. All these spec- Total length to 188 mm (but large mead- imens are from central Wisconsin. Included ow voles seldom exceed 165 mm), tail 42-56 with the value (5) were dark ochraceous No. mm, hind foot 20-23 mm, and ear 20-23 min. 4579 from Waupaca County, 4851 from Ver- Weights vary to 56.9 g. Greatest length of non County, and 5024 from Clark County. skull varies up to 28.6 mm, width to 15.6 From Marathon, Wood and Portage counties mm (see Jackson, 1961). a large sample fell into these frequencies: pure Dental Formula. I 1/1, C 0/0, P 0/0, M silvery white 1; buffy white 16; whitish buff 3/3 = 16. 37; buffy ochraceous 46; dark ochraceous 4. Geographic Distribution. The meadow Such frequencies were similar in other parts vole is found throughout the state, in boreal of Wisconsin. Jackson (1961) almost named and grassy habitats, marshes, bogs, wet and Wisconsin meadow voles as a subspecies char- dry prairies, agricultural land, meadows, acterized by buffy venters. From the foregoing rights-of-way, and even lawns. it follows that Microtus pennsylvanicus can- Status. The meadow vole is abundant, one not always be distinguished from Pitymys of the most abundant mammals in the state. It is ochrogaster by the color of the belly, but white injurious to humankind, especially to farmers, bellies are characteristic of pennsylvanicus. eating grain and the plants themselves; girdling From Portage County, a gray vole had bushes and trees, causing damage in orchards; hairs whitish basally. Jackson (1961) described trenching lawns, golf courses, and cemeteries; yellow voles from Alderly, Dodge County. Two competing with rarer species; and it constitutes albinos from Madison and 6261) is from Hori- something of a health threat by transmitting dis- con Marsh. This vole was normal above ex- eases to other mammals, and by its parasites in- cept for a faint intermixture of white hairs be- directly spreading diseases. There are few mice low the ears and approaching the vibrissae. in captivity as objectionable in aggressiveness and filthiness; special precautions must be employed. To be fair, they are neither likely to bite nor to be filthy in their natural surroundings. They are, by virtue of numbers, very important members of food chains in wetlands and prairies. They constitute a major food item for most raptors, and most of the carnivores. For some of these predators meadow voles are the chief food eaten. Snakes also eat them. The rare great gray owl apparently feeds almost exclusively on this species. Wisconsin’s authority on raptors, the recently deceased Mammalogists also use dental records. Microtus Fran Hamerstrom, found that raptors, and pennsylvanicus, showing posterior islet on middle molar. especially the harrier, depend so much on Pitymys ochrogaster [and other voles] lack it. Synaptomys cooperi, from Door County, shows deep labial (lateral) re- these voles that breeding output is closely re- entrant angles, and a faint, lateral incisor groove. lated to the cycles of the voles.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 281 Habitats. Meadow voles are found on wet, grassy or weedy soils, fields, and wet meadows, in marshes, bogs, along riparian grassy shorelines, in grassy glades or open woods, occasionally in cultivated fields, often occurring on lawns, in gardens, and rarely entering houses. The meadow vole prefers black soils and wet environments due to a need for water. It swims well, even diving, and young have been seen struggling over the water surface (Long, 1970a). Jackson (1961) mistakenly believed that the shrew Blarina brevicauda was the most common mammal in Wisconsin. He suggest-

Maps showing geographic distribution of Microtus pennsylvanicus in Wisconsin and North America.

282 THE WILD MAMMALS OF WISCONSIN ed that if M. pennsylvanicus were found in Foods. This vole eats seeds and green more diverse habitats then it would be the most parts of grasses, sedges, and forbs. The mouse abundant. (I would judge the most common also eats hay and cultivated crops, berries, wild mammal in Wisconsin to be either the insects, fungi, occasionally small bird eggs, meadow vole or the white-footed mouse.) He and small vertebrates, including its own kind. suggested that the meadow vole did not occur The digestive system is somewhat inefficient in sandy areas, dense timber, or gardens. I have (Golley 1960) and these voles occasionally not taken them in dense forest where grasses engage in coprophagy (Oelette and Heising- are absent from the forest floor, for they dwell er 1980). Several examples of cached food in and feed upon forbs and grasses. I have tak- hoards have been found, but there is no com- en them in swamps, bogs, along streams, riv- mon pattern for hoarding food. Several others, on golf courses, lawns and other bluegrass er students on this subject include Zimmer- areas, in gardens, sandy areas, Christmas tree mann (1965), Maxson and Oring (1978) and plantations, and brushy savannas. The great- Gates and Gates (1980). est densities are, indeed, as Jackson (1961) Reproduction. This vole, which often mentioned, in the rank grasses, reed and cat- increases dramatically in a 4-5 year cycle, is tail marshes, meadows, grassy orchards, vines, remarkably fecund. It breeds both early and briars, fencerows, pastures, hay, alfalfa and late in the year (Keller and Krebs 1970), has clover fields, wet prairies, dry prairies where large litters, and has a postpartum estrus. The vegetation is lush, riparian vegetation, railroad annual number of litters may number from 4- and highway rights-of-way and beach grasses 10. In contrast to the generally monogamous (rarely). These voles seem to require runways prairie voles, the meadow vole is promiscu- in grassy or sedge vegetation, where cuttings ous (Getz et a1. 1981) and the female will and runways proclaim their presence. Publica- repeatedly mate (Madison 1980). The females tions treating this wide-spread and abundant are somewhat territorial (Burt 1940). The mammal include Getz 1960, 1961, 1970; Blair young become sexually mature in less than a 1940; Manville, 1949; Golley 1960; Ozoga month (Timm 1975). The gestation period is and Verme 1968; Dice and Sherman 1922; about three weeks. The litter size exceeds 4.5 Hamerstrom 1979, 1986; Mumford and Wh- in several published studies (5.6 in Illinois, itaker 1982; Getz et al. 1981; Krebs 1977; Hoffmeister; 4.54, 5.8; and 4.81 in Indiana, and Tiatt and Krebs 1985. In winter, snow Keller and Krebs, 1970, Mumford and Whi- provides security for the vole as it makes its taker, 1982). The range is 1-9 young. rounds under the snow.. The snow moderates In Wisconsin Microtus pennsylvanicus the microclimate as well as providing insula- can breed every month of the year (Jackson, tion against cold air temperatures (Pruitt 1954). 1961: 233) and usually continues into No- Nests are constructed of grass clippings in vember in central Wisconsin, a full month lat- short underground burrows, in the surface run- er than Peromyscus. In 27 pregnant females way also, under rocks or pieces of wood, even in from Wisconsin, the mean embryo count was the tunnels under the snow. The nests are glob- 5.2, with modes of 4 and 6. The observed ular, with at least one entrance. Some are mere range was 4-8. The peaks of breeding were platforms in the runways. Some are in tunnels in April and July. The females were found 8-9 inches below the surface of the ground. Jack- pregnant from March to October and lactat- son (1961) provides a photograph of such a nest. ing until 14 November. Weilert and Shump (1977) provide dimensions The young weighing about 3 g each are for 20 nests, which averaged 148 mm diameter, born naked and blind (Whitmoyer 1956). In 72 mm depth. Nests are kept clean, which is in five days the incisors erupt and short pelage marked contrast to their cages and live traps. has grown out. The molars erupt by seven days,

TAXONOMIC ACCOUNTS / ORDER RODENTIA 283 and begin to open by eight days. The female stable. According to Burt (1940, 1948), the may be in mid-pregnancy when her previous home range exceeds 1/15 acre (267 sq. m), young are weaned, often by the 12th day. or 1/10 acre (400 sq. m). The known range Young have been live trapped out of the nest is 0.15 acre to 0.75 acre (0.06-0.3 ha) (Blair as small as 10 g, suggesting some young leave 1940; Getz 1961; Manville 1949). Within the the nest even younger than 12 days; the 12- home range there are defended sites, both at day-old voles average 14 g (Golley 1960). Age nests and feeding areas (Ambrose 1973; Burt may be determined by weight of the eye lens- 1943). Females are more territorial than males es (Thomas and Bellis 1980). Sexual maturity (Madison 1980). There is considerable over- is poorly correlated with age because the young lap in home ranges and males wander more may breed early in life. Angularity of the crani- than do females. The habitat affects home um, long rostrum, closure of the basioccipital- range size, as does the availability of food and basisphenoid suture, and body weight all may the density of voles. provide suitable criteria for aging this impor- Diet and predation and the previous year’s tant species. Some recent workers consider the reproductive success greatly affect the local exact age of sexual maturity in these “repro- abundance of this species. Floods doubtless ductive machines” as irrelevant, and use bio- drown many in the wet habitats. Hamerstrom mass instead of “age” for life tables. (1985, 1986) shows how the mouse is not only Mortality. With a high biomass and po- affected by predation but its abundance allows tential energy available to nearly all carnivo- harriers to breed more frequently in a sum- rous species in Wisconsin, this prey species in mer. Being especially abundant in appropri- natural habitats is surprisingly beneficial. It is ate habitat, the interrelations between this vole preyed upon by such diverse predators as game and other mice are important (Grant, 1969; fishes, snapping turtles, probably other turtles, 1971). Meadow voles seem to adversely affect probably bullfrogs in the southern counties, Synaptomys cooperi (Getz, 1961; Linzey many snakes, most owls, most hawks, ravens 1981), where bog lemmings avoid pure grass and crows, herons, shrikes, sand hill cranes, stands, but meadow voles lose out in competi- gulls, short-tailed shrews, dogs, cats, muskrats, tion with the smaller pine voles, (Novak and and most other carnivores. Getz 1969). (The rare pine vole has not been Doran (1954), Jackson (1961), and Timm studied here.) Red-backed voles seldom com- (1985) review the numerous parasites of the pete with Microtus (Iverson and Turner 1972) meadow vole. These include blood protozo- and where found with Peromyscus, the deer ans, cestodes, nematodes, spiny-headed mice may outnumber Microtus as much as 12 worms, and flukes (Zajaac and Williams, 1980). to 1 (Brewer and Reed 1977). Blarina feed Ticks (at least 11 species), mites (more than on meadow voles in summer and winter (Eadie 80 named species), lice, botfly larvae, and fleas 1952). Prairie voles and meadow voles segre- infest these voles (Hatt, 1930b; Mauer and gate into respective microhabitats (Krebs 1977; Skalely 1968; Manville 1949; and Scharf and Zimmermann 1965; Miller 1969). Steward 1980). Parasitic beetles (Leptinidae) Jackson (1961) explained that the peri- also infest this vole on occasion (Timm 1985). odic increases in vole populations are due to Home Range and Density. Home range three factors: accelerated breeding rate, in- is usually defined as the area traversed in nor- creased litter size, and increased number of mal activities. Madison (1985) bases home litters. All of these, of course, would increase range on daily range. He divides the spatial the abundance of a population. Population distribution as stable home range, variable crashes are likely the result of high predation home range, and shifting home range. About and food shortages. Consequent shortage of 3/4 of home ranges in voles are reportedly food for the predators can cause voles to dis-

284 THE WILD MAMMALS OF WISCONSIN perse, lower their breeding performance, or cause voles to starve or die from other causes. Concomitantly, as the predators are on the wane, the voles increase while the food availability naturally increases. This is the concept of seasonality. The vole builds up its population in subsequent years depending on the survivors lasting the winters, and peak numbers cycle up again. Being a complex dynamic system with numerous variables, the cycle may not be the same in all regions, nor may it be in synchrony from one region to Multi-annual Cycle for Microtus pennsylvanicus in another. Basically there is a 4-5 year cycle or central Wisconsin. Nora Lopez-Rivera, from Hamerstrom a 3-4 year cycle (Krebs and Myers, 1974; Tiatt journals. and Krebs 1985). Not surprisingly, there are also annual fluctuations in some places or Green Bay, 1. Florence, Fond du Lac, For- some decades. Jackson (1961) believed that est, Grant, Green, Iowa, Iron, Jackson, Ju- meadow voles over large regions may attain neau, Kenosha, Kewaunee, La Crosse, densities of over 200 per acre, but in Michi- Lafayette, Lincoln, Manitowoc, Marathon, gan Aumann (1965) found densities to be 50- Marinette, Milwaukee, Monroe, Oconto, 60 per acre (21-25 per ha). Oneida, Outagamie, Ozaukee, Pepin, Por- Using 25 years of data collected on the tage, Racine, Richland, Rock, Sauk, Saw- prairies of Waushara and Portage counties by yer, Shawano, Sheboygan, St. Croix, Trem- Fred and Fran Hamerstrom, one of their as- pealeau, Vernon, Vilas, Walworth, Wash- sociates Nora Lopez Rivera found some devi- burn, Washington, Waukesha, Waupaca, ation from a 3-4 year fluctuation. Neverthe- Waushara, Winnebago, Wood counties. less, the cycle was fairly regular and about five years (see Fig.). Remarks. This mostly nocturnal vole is Genus Synaptomys Baird surprising sociable in nature, but almost vi- Bog Lemmings cious in close quarters or unnatural surroundings. It has been observed to stand on its hind Re-entrant angles of cheek teeth deeper on legs and box or threaten much larger ene- one side, shallow on the opposite, so that mies (i.e., human) and even mobbing blue- prisms extend as nearly transverse lophs birds (see Long 1970a). Confined meadow across the tooth surface. The salient angles voles fight and may eat one another. The adults often eat the young when they are found out of the nest. The female is attentive to its young, returning them to the nest with “mother love” (Bailey, see Jackson, 1961). Additional Natural History. A biology of the meadow vole was published by Reich (1981). Geographic Variation. None was observed. Specimens examined. Total 548. Ad- ams, Ashland, Bayfield, Burnett, Clark, Columbia, Crawford, Dane, Dodge, Door Co.: Lost Lake Bog, Chambers Island, in Sketch of Southern Bog Lemming by R. P. Grossenheider.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 285 are reduced on the outerside of the lower gles are exceptionally deep, whereas the in- molars, and on the inner side of the upper ner angles are shallow, so that the upper teeth molars. There is a tiny groove extending along are zagged and not zigzagged. The lower teeth the outer border of the anterior face of each have the deep re-entrant angles on the in- upper incisor. side, and the shallow angles outside, so that these teeth are zigged and not zigzagged. On the middle lower molar is a small outer prism. Synaptomys cooperi Baird The short tail, coarse grizzled fur, Southern Bog Lemming grooved incisors, squarish braincase, and one- sided re-entrant angle pattern distinguish this 1858. Synaptomys cooperi Baird. Report Explor. species. Externally the bog lemming resem- Surv.. Part 1 Mammals, p. 558. The type local- bles Pitymys pinetorum because of its short ity has been fixed at Jackson, New Hampshire. tail. There is a superficial resemblance in their 1896. Synaptomys fatuus Bangs. Proc. Biol. Soc. skulls as well because P. pinetorum has such Washington, 10: 47. Type from Lake Edward, a broad skull. Comparisons of the races are Quebec. given in their accounts below. 1912. Synaptomys cooperi fatuus: Cory. The mam- The upper parts are a mixture of gray, mals of Illinois and Wisconsin. Field Mus. Publ., pale ochraceous brown, and dark brown or 153: 237. black guard hairs. The venter has a whitish, grayish, buffy, or tan ochraceous wash over The name Synaptomys means a mouse dark plumbeous gray bases of the hairs. The that yokes something together. In this case, feet are usually brownish but sometimes gray. the name stands for a mouse that stands evo- The tri-radiate baculum has a stout basal lutionarily intermediate, so to speak, between stalk (Burt 1960). There are eight mammae in voles and true Arctic lemmings (Arvicola and the female. The chromosomes number Lemmus). There is something misleading 2N=50, FN=52 (Hoffmann and Nadler, 1976). about the vernacular name southern bog lem- The size is comparable to that of other ming. Southern bog lemmings often occur in Wisconsin voles, but adults average larger than habitats that are not bogs. Furthermore, this all except Microtus pennsylvanicus. Four species is southern in relation to a northern adult males from Drummond averaged in to- bog lemming, found northward. The south- tal length 113 (110-115), tail length (16 (10- ern bog lemming is boreal, and populations 19), hind foot length 17.6 (17-18), and length found in the south of the range are usually of ear from notch 9 (5-10). In cranial mea- relicts from a cooler time when Pleistocene surements six adults from Gogebic and Tay- glaciations had displaced these mice southward. lor counties, Michigan, and two from Bay- Description. The southern bog lemming field County averaged in condylobasal length is a chunky mouse with tiny tail and grizzled, 25.0 ± 0.23 (24.4-25.5), nasal length 7.05 coarse pelage (except drabbed in worn win- ± 0.05 (7-7.2), zygomatic breadth 15.2 ± ter fur). The tail and short ears are often hid- 0.33 (14.6-15.7), lambdoid breadth 12.25 ± den by pelage. There is usually a faint groove 0.1 (11.9-12.45), and cranial depth with bul- along the anterior face of each upper incisor, lae 8.5 ± 0.13 (8.2-8.8). near the outer border. The skull is subquad- Dental Formula. I 1/1, C 0/0, P 0/0, M rate, i.e., nearly square, resembling Arctic lem- 3/3 = 16. mings in the projections of the braincase out- Geographic Distribution. The southern ward into the orbital space. The rostrum is bog lemming occurs almost always in low short and the zygomata bowed out. In the numbers, but may attain local concentrations. upper cheek teeth the outer re-entrant an- The occurrences are found from one end of

286 THE WILD MAMMALS OF WISCONSIN the state to the other. They are not known in once in great abundance in blue grass near a the Apostle Islands in Lake Superior or in muddy stream. Subsquently he took them in some of the southwestern prairie counties. much smaller numbers. They have been tak- They are found on islands off the Door Pen- en even in beech-hemlock woods (Hamilton, insula in Lake Michigan. See Map. 1941) in New York. In upper Michigan they Status. The southern bog lemmings are are found in spruce bogs, tamarack bogs, and seldom common, and are harmless to human- wet and dry hardwood forests (Dice and Sher- kind. They are important in food chains in man 1922) and blueberry barrens and pine wet-soil habitats and bogs, as well as on black hardwoods (Robinson 1975). One problem soils of highlands. Often found near water, for Synaptomys seems to be the presence of the bog lemmings seem adversely affected by Microtus pennsylvanicus (Getz, 1961). Bea- Microtus pennsylvanicus and by human land sley (1978) found P. ochrogaster and Syn- use. No irruptions have been noted in Wis- aptomys to cycle concurrently in Illinois. Bog consin, as seen in other states. The race on lemmings avoid pure grass habitats and sur- the small isles of Door County, in Lake Mich- vive at brushy edges (Linzey 1981). igan, named S. c. jesseni, has a very limited When bog lemmings are abundant there distribution. For that reason it is quite vulner- are complex systems of runways extending able to eradication by humankind. There is and branching through the sedges and grass- quite a lot of land protected from develop- es. These are littered with grass clippings and ment on Washington and Rock islands, but greenish scats. Sometimes when Synaptomys no effort has been made to protect the sedge are present, but not abundant, there is no sign meadow habitats. The southern bog lemming of runways at all. Burrows are dug in the S. c. gossii, from western Wisconsin, repre- ground, as are tunnels in snow. In some bur- sents a small, localized population. It proba- rows young are reared without nests of grass bly has no protection, because its geographic (Hoffmeister, 1947; Hayase, 1949), but grass range is practically unknown. The southern nests have been reported (Jackson, 1961; bog lemming has declined in Lower Michi- Connor 1959; Burt 1928). Jackson says the gan (Master, 1978). grass nest has up to four entrances and is lined Habitat. The southern bog lemming is with fur, feathers, or fine grasses. occasionally taken in wetlands. I examined a Foods. Conner (1959) concluded that the specimen taken by Russ Mumford, put on dis- chief foods are green monocots, especially play at the Purdue Science Camp, trapped in grasses and sedges. Other foods reported in- the tamarack and sphagnum bogs near the cluded mosses, liverworts, fungi, ferns, forbs, Pine and Popple rivers, in northeastern Wis- woody plants, fruits of raspberry, blackberry, consin. From throughout Wisconsin, Long huckleberry, cranberry, plant roots, bark, in- (1990), Long and Long (1988), Clark (1972) sects, snails and other invertebrates. This vole and Jackson (1961) describe a variety of hab- usually eats the basal and seed parts of a grass itats including sedge meadows, fencerows with or sedge and leaves the stem as a “cutting”. red pines, upland hardwoods, Christmas tree The green fecal scats indicate green vegeta- plantations, marshlands, and bouldery ground tion is important in the diet. in spruce forest near the Rib River. Long Reproduction. Following the usual tem- (1964b) caught one on black bouldery ground poral breeding pattern in Wisconsin mice, near a lake in southwestern Ontario. Timm Synaptomys probably breeds in spring and (1975) caught them in mesic, mixed conifers autumn (April through September). The mice and hardwoods in Minnesota. In Indiana, are so scarce that little is known about their Mumford (1969) caught them in grasses and pattern in Wisconsin. The only evidence for sedges. Hoffmeister (1947, 1989) took them breeding in this collection is from Washing-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 287 Ventral skull of Synaptomys. After Hall.

Maps showing geographic distribution of Synaptomys cooperi in Wisconsin and North America.

288 THE WILD MAMMALS OF WISCONSIN ton Island, an isolated relict population. Bea- The fluctuation of numbers in this spe- sley (1978) found breeding peaks in Illinois, cies is mentioned often in the literature. It but the bog lemming breeds anytime in the seems a “boom or bust”. There are interrela- year according to Burt (1928). Males are re- tions with other voles (see Getz 1961), ef- productive at least from April to December, fects of climate, nutrition, and of course land and females have a postpartum estrus. use. Over-trapping can cause eradication. The Hoffmeister (1989) found breeding pregnan- cycle seems to be a 3-4 year cycle (in south- cies or births in S. cooperi cooperi from April ern Illinois at least) (Beaseley 1978). In south- into November. In Pope County, Illinois, 65 ern Michigan Blair (1948), Linduska (1950), percent of females were pregnant in late and Burt (1940) suggest peaks in 1936, 1938, April. The gestation period is about 23 days 1942, and 1946. Lows were observed in (Connor 1959). Litter size averages about 1937, 1939, 1940, and 1941. three (range 1-7) in Ontario, and in one sam- The occurrences are quite local in dis- ple from Illinois (19 pregnant females) 3.58 tribution in Wisconsin, which can be one rea- (Hoffmeister, 1989). Beasely (1978) listed son why a person might catch 20 bog lem- the mean as about 3 (1-8). mings one year and none the next. Linduska At birth, young southern bog lemmings (1950) caught 18 on half an acre, which weigh about 3.9 g each, and have scant fur makes a very high density of 36 per acre dorsally (Hoffmeister, 1989) or are naked (0.4 kg). Hoffmeister and Warnock (1947) except for vibrissae (Connor, 1959). They obtained 35 in an acre in one year, but in are born blind with ears closed; eyes open most subsequent years the numbers were low in 10-12 days. Weaning begins in some fe- or medium. Usually the distribution is thin- males in 14 days, and seems complete in ner than that, of course; other estimates are three weeks. The juvenal fur is replaced with 14 per acre (Stegeman, 1930), and 2-14 an adult pelage by 60 days. The males may per acre (Banfield 1974). On Washington be sexually mature in 35 days (Connor, Island when bog lemmings seemed abun- 1959) but usually both sexes are breeding dant (they had never been taken previously 60 days after birth. or since) in a hedgerow and lawn (about one Mortality. Synaptomys is taken by diur- acre) six were taken in two nights. Others nal and nocturnal predators. These include were observed on other dates feeding un- snakes, screech, barn, long-eared, great- der a bird feeder. On Swenson Road, bog horned, and even the great gray owls, nu- lemmings were not taken at all except in a merous mammalian carnivores, such as red sedge meadow of about two acres. They did and gray foxes, bobcat and house cat, coy- not range through the meadow, but were ote, gray wolf, probably shrews, all the wea- concentrated at one end. Three were taken sels, mink, and martens. Doran (1954) men- on one occasion, but usually one or none tions tapeworms are common parasites. Con- on other evenings. These data give an esti- nor (1959) and Timm (1975) report mites, mate of about six per acre. sucking lice, fleas, and ticks. These parasites probably contribute to mortality. Home Range and Density. Burt (1940) Synaptomys cooperi cooperi Baird recaptured an adult female seven times in Cooper’s Southern Bog Lemming August and September in Lower Michigan, and obtained an estimate of home range about Synonymy given for species above. 0.5 acre. Buckner (1957) estimated home Description. As defined by Wetzel (1955) the range at 0.11 acre for males and 0.14 acre nominate race for the bog lemming occurs for females in Ontario. both east and west of Lake Michigan. It is

TAXONOMIC ACCOUNTS / ORDER RODENTIA 289 characterized by a rather narrow, high crani- Camp Susan 3. Marathon Co.: 7 1/2 mi. um. All specimens examined in the UWSP NE Athens on Big Rib River 1. Marinette Co.: collection, except those from Door County, 9 mi. NW Crivitz, County A 3. Oconto Co.: proved referable to this subspecies. They show 11 mi. NE Suring 1. Portage Co.: Dewey little variation in color. Some specimens had Marsh 1. Taylor Co.: Near Medford 1. a rich reddish chocolate color intermixed with ochracaeous, gray and black. Winter specimens were bleached and less grizzled. Young Synaptomys cooperi jesseni Long specimens were darker brown. Specimens Jessen’s White-footed Bog Lemming from the Upper Peninsula of Michigan were referable to S. c. cooperi. 1986. Synaptomys cooperi jesseni Long. Mamma- Using a scale from reddish to grayish lia, 51: 324. Type from Ted Jessen’s property brown, the dorsal pelage being lined with dark on Swenson Road, Washington Island, Door guard hairs and darkened from below by the Co., Wisconsin. Named in honor of Tom Jes- underlying basal gray, the most reddish spec- sen, Washington Island, the former Supervisor imen, was No. 5742 from Portage County. of Rock Island State Park. A shade paler is Mus. No. 2197 from Crivitz. Next is 5038 from Bayfield County, where Description. In dorsal and ventral color- the light ochraceous color separates from the ation the specimens from Rock and Wash- dark brown lines of the guard hairs, and final- ington islands, remote and widely separated ly the ordinary coloration—a grayish, ochra- from the mainland bog lemmings (Long and ceous brown, as seen in 5626 from Bayfield Long, 1988), resemble the nominate race. County. Scaled from 4 to 1, all the adult and However they have conspicuous white feet. probably mature bog lemmings were com- The tails and feet are generally grayish, but pared against these specimens taken as stan- there is usually pure white distally in most of dards, and the color values were recorded. In the claws and toes. The toes and claws may the far north, from Fish Hawk Lake, Gogebic occasionally be whitish in S. c. cooperi, but Co. Michigan, the skin was reddish (3). From they are seldom pure white. The whiteness is Menominee County, Upper Michigan, four not nearly as extensive, not set off cleanly by specimens averaged 1.5 (1-2). A vole from dark pelage of the feet, and not nearlv so Delta County, Michigan was 3. constant. White toes are even fairly constant From Bayfield County, five specimens on the forefeet in the insular specimens. averaged 1.7 (1-2), and another with worn Even the hind dew toes are white on pelage (in June) was dark grayish. Langlade the Rock Island specimen. They are white County specimens were 1 and 3, averaging also on UWSP 6548 and 6549 from Wash- 2. An Oconto County specimen was a 2. A ington Island. All ten toes are white on these specimen from Marathon County was 1, and specimens. On Washington Island, eight dis- the Portage County specimen 4. A specimen tinctively white hind toes (excluding dew from Douglas County was 1 (Long, 1990). claws) were seen in each of eleven speci- For measurements, see account of the mens. Seven white toes were seen in the species above. other two, and even in these there was at Geographic Distribution. See Map. least a trace of white in all eight. In 22 adult Status. See account of the species. mainland and Upper Michigan specimens Specimens examined. Total 34. Ashland whitish toes were seen in only five speci- Co.: 6 mi. SE Clam Lake 9 . Bayfield Co.: mens. Some specimens from Menominee Drummond 12. Burnett Co.: Crex Meadows County, Michigan, show several white hind 1. Douglas Co.: Wascott 2. Langlade Co.: toes, as does one specimen from Bayfield

290 THE WILD MAMMALS OF WISCONSIN County, in northwest Wisconsin. But these Synaptomys cooperi gossii (Coues) from the mainland are less distinctive than Goss’ Southern Bog Lemming the least whitish-footed specimens on Wash- ington and Rock islands. 1877. Arvicola (Synaptomys) gossii Coues. In Coues White toes are seldom observed on the and Allen. Monograph N. Amer. Rodentia, p. forefeet of any mainland specimens, but all 225. Type from Neosho Falls, Woodson Co., the front toes were pure white on the Rock Kansas. Island vole, and toes of the front feet were 1897. Synaptomys cooperi gossii: Rhoads. Proc. white on seven from Washington Island. Some Acad. Nat. Sci. Philadelphia, 49: 307. toes of the forefeet were white on the others also, except two of the eight specimens had This bog lemming resembles S. c. coo- only dark toes on the forefeet. White toes peri but is larger and relatively larger in many seem a sporadically appearing character in dimensions. It has especially broad incisors bog lemmings, but on Washington and near- and high cranial depth of the skull. The first by Rock islands it is ordinary (and in fact con- reported Wisconsin specimen, referred to S. stitutes a remarkable and conspicuous differ- c. gossii by Jackson (1961), has wider inci- ence allowing the large majority of bog lem- sors than does cooperi. Across the Mississip- mings to be identified on sight). There is also pi in Minnesota and northeast Iowa, bog lem- a difference in the skull; the island mice have mings also have been referred to S. c. gossii. significantly broader skulls. S. c. gossii occurs on hillsides and fields Jessen’s bog lemmings from Washington in southern and southwest Wisconsin. The Island, Wisconsin, averaged for total length in habitats seem to be dryer than usual for bog six males and three females, respectively, 118 lemmings. They may be expected to occur in (112-122), 123 (119-126), tail length 17 (15- wetlands as well. This race was named in 19), 18 (16-20), hind foot length 19.4 (18- honor of an early Kansan naturalist B. F. Goss. 21), 19 (17-20), and length of ear 11 (10-12), Recent specimens from Fort McCoy are 11.3 (11-12). Six adults from Washington Is- assigned to this race, not so much by size, but land and one from Rock Island averaged 23.6 the skull seems deep in cranial depth, the inci- ± 0.2 in condylobasal length, 6.87 ± 0.14 (6.3- sors are notably wide even in young bog lem- 7.3) in nasal length, 15.67 ± 0.03 (15.6-15.8) mings, and the venter was invariably rusty ochra- in zygomatic breadth, 12.93 ± 0.13 (12.6- ceous (whereas in S. c. cooperi the venter is 13.4), and cranial depth including bullae 8.62 usually whitish, never this brightly ochraceous ± 0.1 (8.3-9). or tawny). These bog lemmings were taken on Geographic Distribution. Rock and Wash- highlands in oak savanna and grasses, as was ington islands, Door County, Wisconsin. Jackson’s record from Lynxville. A female was Status. See account of the species. This lactating, a male was in breeding condition, and race has a very limited distribution, on two young animals were taken 28 August 1995. small islands. Specimens examined. Total, 5. Crawford Remarks. Cuttings of sedges in the mouth Co.: Lynxville, 1. Monroe Co.: Fort McCoy, 4. of UWSP 6259 were 3, 4, 5, 5, 6, and 8 mm in length. Pregnant females were taken in April (5 embryos) and late September (3 embryos). Genus Ondatra Link Specimens examined. Total 14. Door Muskrats County: Rock Island Hq., 1. Washington Island, Swenson Road, 6; Airport on Airport Road, 7. The muskrat was first named by Linnaeus, The holotype and a topotype are deposited in who considered it a kind of beaver. To this the U. S. National Museum (Nat. Hist.). day many mammalogists consider it a web-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 291 footed beaver-like creature, but it is not. The Russia, China, and much of Europe. One dif- hindfoot at best might be regarded as par- ference between Ondatra and the Wisconsin tially webbed. This brownish, long-tailed, voles, aside from much larger size and the aquatic mammal is more exactly a gigantic, aquatic specializations, is the presence of five long-tailed vole than a beaver. It has the pe- functional clawed toes on all four feet in the culiar zig-zag molar teeth of voles. The mo- muskrat (the thumb bears only a small nail). lars show salient and re-entrant angles, comprising in the occlusal view a series of prism- like islands of dentine more or less enclosed Ondatra zibethicus (Linnaeus) by sharp-angled walls of enamel. The aquatic Ondatra zibethicus zibethicus (Linnaeus) specializations include the following: the hind Muskrat feet have conspicuous fringes of stiff hairs, and the long scaly tail, compressed laterally “What makes the muskrat work so hard to guard its for sculling propulsion, has a small fringe. The musk? Courage!” — Lion in Wizard of Oz. fur is not remarkable except that the guard hairs and dense underfur are soft and pretty 1766. [Castor] zibethicus Linnaeus. Systema naturae and also durable. Closely related to Neofiber, Ed. 12, 1: 79. Type locality of the nominate the round-tailed muskrat of Florida, this North race and the species is Eastern Canada. American genus contains according to some 1795. [Ondatra] zibethicus: Link: Beytraege zur workers a weakly differentiated Newfoundland Naturgeschichte, 1: 76. species, 0. obscurus (Bangs), as well as the wide-spread species 0. zibethicus (Linnaeus). The name Ondatra has no known The latter is the type of the genus. It ranges meaning; probably it was taken from an In- from Alaska through most of Canada and the dian name. The specific part of the binomen, United States, and southward to northern zibethicus, is Latin for musky odor. There Mexico. (The author was with Dr. Syd Ander- is immense confusion about spelling of this son’s field party that collected the first one in name; those who make themselves a rule that a turtle trap in Chihuahua.) Introduced into when the generic name ends with “a” then the Old World, they are now taken for fur in the trivial name also ends with “a” are in this instance led astray. The word zibethicus agrees with Ondatra in gender and number, and should not be revised. An early day corruption of the species name Muskrat to “mushrat’ reminds me of a long ago use in Wisconsin of a binomen “Mus rattus”. Seemingly a silly mistake, this binomen is actually available for the black rat ( = roof rat) Rattus rattus. Description. A cat-sized or smaller rodent having brown fur, elongate, compressed, mostly naked tail (having numerous scales evident), and feet with fringes to facilitate swimming. The skull is similar in form to those in the voles, with the braincase subquadrate and sharply angled, even protruding into the orbits in dorsal view. The rostrum is long and Muskrat. By Alan Long. narrow with flaring zygomata.

292 THE WILD MAMMALS OF WISCONSIN hairs are blackish. The venter is much lighter, tending to a pale grayish ground color overlain with golden, cinnamon, or tan col- or, occasionally reddish tawny, and sometimes tan or gray-fawn brown. The mid-dorsum and legs may approach black because of the intermixed black hairs. The head is concolor with the body. The short ears are practically buried in the fur. The tail is slate gray or darker, sometimes brown, and the naked parts of the feet are much the same in color, gray or brown. Jackson (1961: 246) reported two Skull of Ondatra zibethicus. albinos from Wisconsin. Occasionally there are yellowish or cream colored mutants. The interorbital breadth in comparison Exceptionally large for an arvicoline ro- to the zygomatic breadth is relatively narrow- dent, the adult muskrat exceeds 400 mm, the er than in the voles. The dentary is very sharp- hind foot is greater than 60 mm, and the adult ly bent, with the lower incisor and its root skull exceeds 55 mm in greatest length. nearly straight and having a sharp angle with Weights vary from 2-3 pounds (1-1.4 kg). the axis of the posterior part of the dentary. Dental formula. I 1/1, C 0/0, P 0/0, M The incisors are usually orange and quite for- 3/3 = 16. midable. In occlusal pattern the cheek (mo- Geographic Distribution. The muskrat is lar) teeth are zig-zag with flat surfaces. The found in boreal forests, prairies, wetlands, large upper third molar’s pattern is distinc- streams and lakes, along Lake Michigan and tive for there are two or three closed trian- Lake Superior shores, in urban areas, in the gles between the anterior and posterior loops. southern riparian lowland forests, along the The three-pronged baculum is protrusible Mississippi and other huge western rivers (i.e., in adult males. The annual molt begins in au- Wisconsin, St. Croix, and the Chippewa riv- tumn. Adult pelts show mottled molt areas ers). It ranges from the security of the waters and juveniles show bilateral molt areas (Ap- to almost any habitat in Wisconsin during plegate and Predmore 1947, Shanks 1948, autumn dispersal, spring breeding dispersal, Linde 1963). This is the best aging character summer drought, or winter hunger. Along the for the genus, because the teeth are all root- shores of the Great Lakes the powerful wave ed and show no patterns of wear. Teats on a action destroys aquatic emergent and sub- pelt indicate female sex. There are two pectoral and four inguinal mammae. Perineal scent glands are at the base of the tail in breeding adults (Jackson, 1961). I (1978a) discovered a small epiphyseal attachment on the dorsal aspect of the scapula, which becomes fused in adults to the scapula itself. The diploid number and fundamental number of chromosomes are 54 (Hsu and Benirschke, 1971). Muskrat fur is usually a dark, rich walnut brown with tones of dark rust, cinnamon, or Summer lodge of Muskrat. A. MacKinnon, age 13, from a golden-bronze sheen. The intermixed guard Anna B. Comstock’s Handbook 1911.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 293 merged vegetation, and muskrats are transient in these habitats. They do persist in protected bays and behind sand spits where vegetation can grow. Status. The muskrat is an abundant mammal in Wisconsin. For years this was the most important furbearer in the wild, both in numbers taken and in economic yield from the marketed fur (Boutin and Birkenholz 1987). This mammal is protected by state game laws that regulate the harvest. It benefits from wetlands set aside for fisheries and duck reproduction. The current decline of the fur

Maps showing geographic distribution of Ondatra zibethicus in Wisconsin and North America.

294 THE WILD MAMMALS OF WISCONSIN trade may increase muskrat numbers; there prior to the harvest. In Wisconsin, Harold is recent evidence of an upsurge in the mar- Mathiak (1966) studied Wisconsin muskrats ket (Table Rod-9). for years, and others have kept up the work. Muskrats are usually the most trapped of Errington (1963) in Iowa conducted a series the furbearers in Wisconsin because of their of masterful studies. abundance. Fur prices are high enough to Habitats. Some of the various and diverse create a demand and increase the trappers. habitats where muskrats may be found are Aside from fur this mammal makes little eco- detailed above. The preferred habitat of this nomic impact. The flesh is edible, and occa- species is one kind or another of wetland. sionally is eaten. Some Native Americans have Additionally, there must be available vegeta- regularly eaten them. Occasionally they are tion which can be used for food, and in many sold in food markets in other states. Muskrats places in Wisconsin this vegetation is essen- damage dikes (Cook 1957) causing trouble in tial to build conspicuous grassy and mud lodg- game management areas. Craven (1984) re- es for winter survival. Ecological requirements ported gnawing to a vehicle and rubber fit- are reviewed by Boutin and Birkenholz (1987). tings on appliances near water. Muskrats may Muskrats may live in burrows excavat- adversely impact native plant communities ed in the banks of ponds, rivers, and other (Willner et al., 1980). Many animals make use bodies of water. They also build houses typ- of their abandoned houses. ically smaller than beaver lodges, but some Muskrats rarely eat crops (see Foods be- are immense. The muskrat house is built of low). Their predatory impact is negligible, but available marsh vegetation, including marsh they do eat mollusks and perhaps other small grasses, bulrushes, and cattails. At the base vertebrates. Integral to the food webs for these houses may be three meters across. marsh communities, they are killed and eat- Jackson (1961) reported a house from the en by some of the larger carnivores, such as Kickapoo River which extended 41 inches wolves, coyotes, and bears, as well as an oc- (1.1 m) high above the ice, and measured casional raptor. 31 feet (10 m) in circumference. The en- Economically valuable, this species has lit- trances are usually below the water level and tle effect on humankind except as a fur resource lead inward to a dry nest chamber above and an indirect value as a wetlands prey spe- the water level. Jackson (1961) sketched the cies. In 1982-3 a total of 7.4 million muskrats interior of a house from Rock County, and were harvested for fur in North America. The a bank burrow observed in Bayfield Coun- $28 million was second only to value for rac- ty. Cory (1912) sketched a bank burrow. coons, which are in fashion (1999). Neither mentioned any nest material inside It is important to learn muskrat biology burrow or house. The natal den reportedly not only because of the economic value of has a nest of grasses for bringing forth the fur harvest, and its consequential econom- young (Smith 1938). ic value in the work place and marketplace, Muskrats in Wisconsin make use of air but also because wetlands are important to trapped under winter ice, but also maintain sportsmen, fishermen and duck hunters, deer breathing holes where they often emerge to hunters and campers. Related to this, is the eat green vegetation. Sometimes so much aesthetic and functional role of the muskrat aquatic vegetation is heaped around the air in the wetland communities. To manage such hole, the muskrat has created an outlier house animals, one must understand fecundity (see in which it eats and rests. In February 1999, Reproduction below), mortality (see below), I observed near my home two whitetails and attempt to understand population biolo- (Odocoileus virginianus) walk on the ice ap- gy. A game manager must know the density proximately 60 m (64 yards) away from shore

TAXONOMIC ACCOUNTS / ORDER RODENTIA 295 to eat the green vegetation thrown out by a Muskrats exemplify Lord’s ecological rule muskrat. Crows also rummage through the for non-hibernating small mammals, that lit- muskrat’s ice-hole vegetation. ter size is significantly larger in northern lati- Foods. Jackson (1961) and Perry (1982) tudes (Danell 1978, Boutin and Birkenholz, list numerous species of aquatic plants on 1987). In Wisconsin, the litters appear from which Ondatra feeds. Cattails and bulrushes late April to August, and there are usually two make up about 80 percent of the diet accord- litters (Mathiak 1966). ing to O’Neal (1949). Occasionally muskrats The young are born blind with a short, range away from the water to feed on corn, fine and dark fur, but with naked feet (Smith soybeans, and alfalfa (Errington 1963). They 1938). Each is about 4 inches in length (about also eat mollusks (mostly clams), dead fish, 100 mm) and weighs about 22 g. At five days and other flesh (Schwartz and Schwartz 1959, they cling to the mother’s nipples, much as Harold Mathiak, personal comm.). Muskrats in some voles, and they may hang on even if also eat algae. On August 6, 1999, Tyler, she enters the water. The eyes open in about Stephanie, Claudine Long, and I observed, in 2 weeks. The fur becomes wooly and gray. Jackson Harbor, Washington Island, a musk- The young are weaned in about 30 days, and rat swimming out from beneath a mat of veg- by 200 days they are adult sized. etation (washed up against the shore). It was Mortality. Errington (1963) suggested carrying a huge bundle of green alga resem- there are two important factors that may dec- bling Spirogyra. The muskrat swam to a imate huge musksrat populations (aside from moored boat and climbed up onto the pro- human caused catastrophes, such as draining peller axle a couple inches below the water a marsh). These were drought (falling water surface. There, about 3 m offshore, it ate the levels) and disease. He believed the popula- entire mass. This alga was bright green and tions compensated for positive or negative showed no sign of gas bubbles. Then the factors by “intrinsic sensitivity” to density, muskrat dived into the crystal clear water, which is in my opinion a meaningless phrase. swam down to the bottom, about ten feet O’Neal (1949) argued for food supply. Fluc- depth, and scooped up another mass of alga tuation seems to me a typical irregular cycle from the bottom. The algal masses were car- due to various factors of mortality and natali- ried with the forefeet while swimming with ty. However, cannibalism and strife in high- the tail and hind feet. The muskrat swam un- density populations (Errington 1940, Mathi- der water out of view. ak 1966) might be a population control mech- Reproduction. Reproduction may occur anism. Adjusted litter size (from 6.35 to 8.41) in any month in the southern states, but seems could naturally regulate population control. limited to the growing season (spring to au- These factors are in need of additional study. tumn) in Wisconsin and Michigan. There may Smith (1938) and Errington (1939) studied be several litters in the season. There is a mortality factors as well. These included se- postpartum estrus. Precocial breeding is rare vere winter frost, flooding, disease, and over- (Mathiak 1966). Muskrats court, with agonis- trapping. In a chaotic cycle an odd thing such tic behavior between males and between fe- as a peculiar predator (such as a crow) might males. Beer (1950) suggested they were mo- trigger unforeseen scaling and irregular oscil- nogamous. Gestation is about 25 to 35 days, lations. Chaos theory has not been applied, and the litters may number from one to 11 nor will it be easy. offspring. Mathiak (1966) found pregnant fe- One of my former students, Makio Su- males in Horicon Marsh from April to Novem- zuki, trailed coyotes across marshes in cen- ber 3; a litter on April 16 suggests breeding tral Wisconsin, and found where coyotes had about mid-March in the southern counties. killed several muskrats. The aquatic mink

296 THE WILD MAMMALS OF WISCONSIN (Neovison vison) is a natural enemy of musk- management studies. Errington (1943) multi- rats (Errington 1943). Other carnivores and plied the number of known breeding territo- some birds of prey also feed on muskrats. ries by the number of young produced by the Coyotes, foxes, bobcats, mink, wolves, dogs, mother. Comparison of placental scar counts and other carnivores may kill and eat musk- to the proportion of juveniles in the autumn rats. Herons (Ardea) and raptors kill them. harvest does not account for mortality of the Humans trap them in great numbers. Many mothers, and autumn samples may not rep- are killed by automobiles. Diseases and resent the total population (Boutin and Birken- drought (with lowered water levels) wipe out holz 1987). Densities may vary from none in entire populations (Errington 1961) and cold what seems good habitat, to about three per frost drives them helpless from their lodges acre. High densities may reach in good habi- to range over the snow and ice, to probably tat as many as 35 muskrats per acre (Banfield fall victim to hungry carnivores. Territorial 1974, Lay 1945). fighting also drives the losers into danger (Err- The fecundity for a Wisconsin female is ington 1939). about 14-15 young per year. About 10-36 Blood flukes, tapeworms, and nematodes percent of autumn born juveniles survived. In are internal parasites (Doran 1954) and mites a follow up study, 18 percent survived (Dom- infest the fur (Smith 1938). Diseases such as ey and Rusch, 1953). In Minnesota and Iowa Errington’s disease, tularemia, and coccidio- the survival rates were 50-53 percent. This is sis cause mortality. an odd difference, which needs to be explained. Home Range and Densities. Home Mathiak (1966) estimated that 87 percent of range, a vague geometric in the best of cir- the muskrat population dies during the first cumstances, is for the general wanderings of year, and 98 percent do not survive two years. muskrats dependent upon water levels, food This is strong mortality for animals that can availability, high versus low density, and land live potentially as long as 10 years (Johnson use by humankind. Home range for muskrats 1925). Obviously juvenile mortality is heavy is determined from recapture data, shoreline (see Boutin and Birkenholz 1987). occupancy, and radio-tagged movements. Muskrats may cycle with annual and mul- Home range was calculated as areas 7-30 m tiannual peaks. In some places there may be a in diameter by Takos (1944). Errington (1963) tendency to cycle every 10-14 years (Errington recorded the diameter as 61 m in Iowa. Mac- 1963, Elton and Nicholson 1942, O’Neal Arthur (1978) studied radio-tagged muskrats 1949, Keith 1963). This cycle may be seen in and found them to wander 5-10 m from the fur returns for the Mackenzie River District of lodge when the water around it is covered Canada (Elton and Nicholson 1942). with ice. Boutin and Birkinholz (1987) report There is some territorial fighting among values ranging up to 30 m from the dwelling, muskrats, especially when the populations are i.e., up to 60 m diameter. high (Sather 1958). The mother will defend Various ways to estimate density are her “territory” when offspring are present. counting houses and using the value for litter Major dispersals occur in spring to breeding size (see Boutin and Birkenholz 1987). None sites. Males and young move about with the of the ways is perfectly reliable. House counts thawing of ice and consequent high water do not account for muskrats in bank burrows, (Sprugel 1951). Sprugel refers to this as the but in marshes there may be no bank bur- “spring shuffle.” Other dispersal is ordinary, rows. The Wisconsin Department of Natural by young-of-the-year, especially in autumn. Resources usually estimate, in marshes, five When water freezes so thick in winter that veg- muskrats per house. Wisconsin’s Horicon etation is cut off from muskrats, they must leave Marsh Fur Farm was established for muskrat their lodges (Mathiak 1966, Sather 1958).

TAXONOMIC ACCOUNTS / ORDER RODENTIA 297 These dispersals have been reviewed by Bou- Subfamily MURINAE Gray tin and Birkenholz (1987). Wandering musk- Introduced Rats and Mice rats may take temporary quarters in burrows of other mammals. Maximum movements are The ancestors of these introduced pests in 180 m in summer (Takos 1944), in winter sel- North America are from Asia, but from there dom more than 10 m. When populations are these mammals had spread over the earth at high density, the young-of-the-year are dis- wherever people dwell. Especially in ports and persing, and due to food shortages, drought, coastal cities the rats and mice arrive in ships or frost the muskrats may move as far as 20 and cargoes, and they eventually find their miles ( = 32 km) (Errington 1939)! way inland. They may be expected anywhere Remarks. Interesting behavior of musk- people have cities, towns, settlements, and rats includes the deposition of scent and fe- barns. They are responsible for many diseas- cal scats at certain sites, probably for territo- es transmitted to humankind directly or rial marking. Muskrats are both diurnal and through parasites (including the infamous nocturnal, with activity peaks in the after- black plague). They greatly damage interiors noon and at twilight (Smith 1938, Boutin and furnishings of homes and apartments, kill and Birkenholz 1987). Muskrats seem anti- poultry, and consume our crops in the fields, social, except in winter lodges or in family barns, and granaries. In slums, they actually groups. Even in those circumstances aggres- attack children and house pets. sion may be expressed when density is high The Wisconsin murines differ in size but (Errington 1940, but see Errington 1939). show resemblance in their drab pelage, In winter there may be large aggregations in brown-gray above intermixed with some black- a single lodge, with up to 20 muskrats ish hairs. The venter is paler than the back, present (Bailey 1937; Lay 1945). but hardly ever whitish. The skull shows su- I once watched a muskrat defeat a mink praorbital, longitudinal but inconspicuous ridg- in a fight, and after winning the muskrat ran es (which are seen in a few of the native mice up and down the bank like a crazed person, in North America too). The murines (Norway “looking for more worlds to conquer”. (I rat and house mouse) have tuberculate (i.e., stayed out of its way!) One in a frozen marsh bunodont) molar grinding surfaces, the trans- caught out in the open was leaping at me, a verse border of the palate lies behind the third threat and not all bluff. molars, and the tails are long, scaly and sparse- Additional Natural History. Willner et al. ly haired. The eyes are small relative to the (1980) wrote a fine review of muskrat biology. head in the rats Rattus. Geographic Variation. There is only one subspecies in Wisconsin. Specimens examined. Total, 74. Adams, Key to Introduced Rats and Mice Brown, Chippewa, Columbia, Dane, Door, Fond du Lac, Forest, Grant, Green, Jeffer- 1 Size small, total length less than 250 mm, son, Juneau, Kenosha, La Crosse, Lincoln, and tail less than 110, occipitonasal length Manitowoc, Marathon, Marinette, Milwau- less than 35, posterior molars quite small, kee, Oconto, Oneida, Portage, Racine, their lengths combined shorter than the Rock, Rusk, Shawano, Sheboygan, Taylor, length of the upper first molar, the inci- Trempealeau, Vernon, Waukesha, Wau- sor tends to spald off behind the anterior paca, Waushara, Wood counties. surface, so that the tooth seems notched Other records (After Balliett and Taft, in lateral view ...... House Mouse 1978): Monroe Co., Ozaukee Co. (No spe- Mus musculus cific localities, not plotted).

298 THE WILD MAMMALS OF WISCONSIN 1’ Size medium, larger than mice with total from human habitations, especially in north- length greater than 250, tail longer than ern and central Wisconsin where winters are 110, occipitonasal length longer than 35 severe. The species often turns up as prey in mm, posterior molars in tandom length the pellets of great horned owls and other longer than length of upper first molar, wild predators. incisors never notched in lateral view .. Several races of Rattus norvegicus have ...... Rattus been introduced into North America, but for 2 Tail longer than half the total length, tem- all practical purposes there is one variable poral or parietal ridges bowed out, 13 kind, with no geographic races apparent. mm or more .... Roof Rat (= Black Rat) Rattus rattus So far unknown in Wisconsin Rattus norvegicus (Berkenhout) 2' Tail shorter than half the total length, Norway Rat temporal ridges nearly parallel, less than 13 mm separation ...... Norway Rat “Ernst Mayr tells of a steamer wrecked off Lord Rattus norvegicus Howe Island, east of Australia, in 1918. Its rats swam ashore. In two years they had so nearly exterminated the native birds that an Islander wrote, Genus Rattus Fischer ‘This paradise of birds has become a wilderness, Norway and Black Rats and the quiet of death reigns where all was melody’.” — Rachel L. Carson, The Sea Around Although the Black Rat or Roof Rat, Rattus Us, 1950. rattus, is not known from Wisconsin, it often shows itself in port cities, such as nearby 1769. Mus norvegicus Berkenhout. Outlines of the Chicago. A melanistic rat is not necessarily Natural History of Great Britain and Ireland. the black rat, and most black rats are not black. 1:5. Type probably from England, assumed to The common Norway rat is seldom seen far be Norway. 1912. Epimys norvegicus (Erxleben): Cory. Field Mus. Nat.Hist., 11: 180. 1916. Rattus norvegicus: Hollister. Proc. Biol. Soc. Washington, 29:126.

The generic name Rattus is a Latinized old English word meaning rat, and norvegicus is possibly a sarcastic joke about Norway as the presumed locality whence the rats came. This inappropriate scientific name is valid. Perhaps this rat, being such an enemy of humankind, should be called a “Dirty Rat,” an epithet made famous by the tough-guy actor James Cagney. Description. See Key characters above, and accompanying figure. The pelage is grayish or brownish except for color aberrations, and the tail is sparsely haired and scaly. The Dorsal aspect of skulls of Rattus norvegicus (left) and R. rostrum of the skull is elongate, the diastema rattus and its upper cheek teeth. After Hoffmeister, 1989. nearly twice the length of the maxillary tooth-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 299 row. Pronounced parietal ridges are widely six feet ( = 1.8 m). It often gnaws holes in separate, and the molars have three rows of wood for ready passage, such as the famous cusps on unworn teeth. The baculum is a rat-holes described in Huckleberry Finn by small, jointed rod. There are six pairs of mam- Mark Twain. The rat runs about under floors, mae. The chromosomes number 2N = 42 tunneling into stored crops and so forth. It (Hsu and Benirschke 1967). A long-tailed follows certain defined paths, which are variant from Portage County has teeth char- called runways. The nests are bulky, made acteristic of R. norvegicus, not R. rattus. of grasses, sticks, and other available mate- Measurements reveal that Norway rats, rials (Baker 1983). contrary to “big-city” myths, are never excep- Foods. The Norway rat eats most any- tionally large (total length 300-482 mm; tail thing edible, especially grain around farms, and 120-215, hind foot 31-44, ear length 15-22 young domestic fowl, garbage, and carrion. mm, and weights vary from 170-850 g, with Reproduction. Breeding can take place the average about 300 g. at any time but is more frequent when food Young rats are difficult to identify, being conditions are good. Females can breed when as gray and small as other rodents; their tail four months old, and may produce six to 12 is scaly and hind feet (which grow faster than litters per year. Litter size is usually 6-8 off- other body dimensions) huge. spring, but Burt mentioned a record of 22 Dental Formula. I 1/1, C 0/0, P 0/0, M (Burt 1946). Gestation is three weeks. There 3/3 = 16. is a postpartum estrus. The newborn young Geographic Range. The Norway rat may weigh about 5-6 g each, and are born naked. occur anywhere in Wisconsin where people In 9-15 days the eyes open. Fur appears by are present. Rats are especially abundant in seven days. Weaning takes place in a month, city buildings, near waterways and run-down when the young are about 45 g in weight. buildings, and in barns and barn lots. Mortality. Enemies of the Norway rat Status. Occasionally locally abundant, the are listed by Lantz (1910) and Baker (1983). Norway rat is injurious to humankind, destroy- Parasites are exceptionally important causing fowl, grain, and property and spreading es of mortality (Doran 1954, Banks 1910, diseases. On the positive side of the ledger, it Jackson 1961, Stiles and Crane 1910) af- eats some weed seeds, and is fed upon by fecting both the rat but also its hosts (includ- owls and occasionally by other predators. ing humans). The albino Norway Rat, with pretty pink Home Range and Density. Home range eyes, is an important laboratory animal, where is rather small, maximum distances travelled it often seems surly and aggressive, if not about 150 feet (Davis et al. 1948). In Balti- downright vicious. It is often kept as a pet, more, the density in the city was 25 to 150 in and I knew a lady who adopted and babied a a city block. A colony usually is comprised of real biting “devil,” loved it, and it became 10-12 individuals. A litter of half-grown young docile and affectionate. Even love-reared rats swells this number considerably. Baker (1983) may cause an allergy in many who handle suggests a high of 300 rats at a single farm. them. They are used as food for pythons and Remarks. These rats form small colonies, boa constrictors, are excellent dissection an- usually eight or ten, and although fighting is imals for general zoology, and serve human- commonplace there are sociable aggregations kind in diverse scientific experiments. almost everywhere the rats thrive. Dominance Habitat. See Distribution above. This rat and submissiveness are discussed by Calhoun makes burrows (2-3 inches in diameter = 51- (1962). They are nocturnal mostly, but can 76 mm), less than 18 inches (= 457 mm) be observed often during the day. Dominant deep in the ground, and varying in length to males seem to suppress breeding in a colony,

300 THE WILD MAMMALS OF WISCONSIN for when the male is removed, promiscuous Mus musculus Linnaeus breeding takes place. House Mouse Additional Natural History. Calhoun 1962; Bjornson, Pratt and Littig 1969; Zins- 1758. Mus musculus Linnaeus. Systema naturae, ed. ser 1935; Henderson and Craig 1932. 10. 1:59. Type from Sweden. Geographic variation. None is evident. Specimens examined. Total, 29. Clark, The name Mus means mouse, and in Old Dane, Fond du Lac, Jackson, Manitowoc, English ‘thief’. The trivial name musculus Marinette, Milwaukee, Oconto, Portage (in- means little mouse. This mammal is also called cludes Rosholt, long tail, but teeth norvegi- the “laboratory mouse”. cus), Price, Racine, Trempealeau, Vernon, Description. See characters in the Key Waupaca, Waushara, Winnebago, Wood above. The house mouse resembles Rattus counties. and superficially, because of small size, it re-

Map showing geographic distribution of Rattus norvegicus in Wisconsin. Known throughout North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 301 sembles deer mice, harvest mice, and jump- longer hind feet. Its skull resembles that of ing mice. The house mouse differs from deer Mus, even in size, but in jumping mice the mice in having darker underparts and a scaly incisors are grooved, and the incisive foram- tail. The skull has tuberculate (i.e., bunodont) ina are much wider. There are eight to ten teeth, the posterior margin of the palate ex- mammae. The diploid (2N) chromosome num- tends farther posteriorly. The incisors are ber is 40 (Hsu and Bernirschke 1967). Prob- usually notched in lateral view; for the pos- ably it is their anal scent glands that produce terior and distal part spalds off. From the har- a musky odor, possibly the urine adds to it, vest mouse Reithrodontomys, the house and there is a mild odor produced by the feet. mouse has a dingier venter, and the tail is For cranial measurements, see Table Rod-14. longer and scaly. There are no grooves on The total length varies from 125-200 mm, the upper incisors in Mus. The jumping mouse and the tail from 60-100 mm. House mice has a white belly, a much longer tail, and much weigh up to 28 g.

Maps showing geographic distribution of Mus musculus in Wisconsin. Known throughout North America.

302 THE WILD MAMMALS OF WISCONSIN Dental formula. I 1/1, C 0/0, P 0/0, M Mortality. The same predators that eat 3/3 = 16. deer mice will also, on occasion, eat house Geographic range and Status. State-wide mice. A great enemy of this mouse is human- and injurious to humankind. kind, appropriately in this case, as well as Habitat. The house mouse in summer house cats and dogs. Parasites are similar to may be found in human habitations but often those of the Norway rat, and the same refer- ranges into the nearby fields, almost any kind ences are useful. Marsh and Howard (1976) of field where vegetation is dense or the food have published an excellent control book. grains are available. In winter the mouse gen- Home Range and Density. Fitch (1958) erally moves into the habitations of human- found Kansas house mice to wander in areas kind, although some probably pass the win- 20-100 feet in diameter. Populations fluctu- ter in the sheltered southwestern coulees and ate significantly, and sometimes there are southern prairies. explosions that make national headlines Nests are made of grass, or in houses of (82,000 per acre) (Hall 1927). Lowery (1974) shredded paper, feathers, or any fine found 500 per acre. Outdoor numbers peak material, in a sheltered place. Baker (1983) in autumn. reports that in fields the house mouse exca- Additional Natural History. Biology of the vates a burrow and makes a nest of grass and laboratory mouse, by Snell 1943. Mouse ge- fine materials for rearing young. netics are summarized by Grueneberg 1943. Foods. House mice eat almost anything, Geographic Variation. None has been preferably seeds, fruits, garbage, and insects. observed in Wisconsin. The mice are a hodge- Reproduction. The house mouse is poly- podge of mutants, their ancestors introduced estrous, has large litter size, and a postpar- from around the world. Possibly they may tum estrus, this mouse has incredible fecun- adapt to certain areas (if not regions) by rapid dity. Females may breed at 45 days age. Ges- evolution. The great majority of the specimens tation is three weeks or even a little less. collected are wild type color (termed “agouti” Young are born pink and naked, with eyes in mouse genetics), but mutants abound. It is and ears closed. They weigh about 1 g each. impossible to assign this variable mouse to Litter size is about 6 (3-13). They are furred any known geographic race. by 10 days. The eyes open by 14. Weaning Specimens examined. Total, 102. Clark, takes place in three weeks, and the female Columbia, Dane, Door, Eau Claire, Green, may already have produced another litter. The Iowa, Jackson, Jefferson, Juneau, Lafay- mice may live a year in houses, perhaps only ette, Lincoln, Manitowoc, Marathon, Mari- a few months in the fields. nette, Milwaukee, Monroe, Oconto, Onei- da, Portage, Price, Racine, Richland, Rock, St. Croix, Trempealeau, Vernon, Waupaca, Waushara, Winnebago, Wood counties.

Table Rod-14. Cranial measurements Mus musculus. Darlington, 5 ad males. Lafayette Co. 6 ad females.

Occip. nasal l. Zygo. Br. L. nasal Max. t-r 21.6 11.25 7.4 3.48 ±1.15 ± 0.5 ±0.4 ±0.15 Palatal views of the skulls of the harvest mouse, house 21.36 11.38 7.56 3.6 mouse, and jumping mouse. After E. R. Hall and his artists. ±1.36 ±0.5 ±0.28 ±0.2

TAXONOMIC ACCOUNTS / ORDER RODENTIA 303 Family ZAPODIDAE Coues Usually (except in Napaeozapus) there is Jumping Mice and Siberian Birch Mice an upper premolar present on each side, conspicuously reduced in size, and the or- Although Klingener (1984) assigned the birch ange or dark yellowish upper incisors are mouse and jumping mice to the jerboas, Di- grooved on their anterior faces. Zapodids podidae, I am unconvinced. Not only is the have much larger infraorbital foramina than Zapodidae a distinctive and ancient group do many murids, including New World (from the Upper Oligocene, mid Pliocene forms of the Muridae. Unlike murids, the epochs), but the Dipodidae shows extreme ovate incisive foramina are tremendously specialization in evolution of a fused tri-radi- wide and deep, probably functional as orate cannon bone in the hind foot and having gans of Jacobson (enhancing taste). The auditory bulla large and some species pos- much folded molars, presence of upper sess a complex “honey comb” inner struc- premolars, elongate hind feet and tail, and ture. Besides remarkable specializations for wide incisive foramina set this group of jumping, some dipodids have fusion of the mice apart from other Wisconsin mice. The last five cervical vertebrae and some even of adaptation of hibernation, lasting in Zapus the axis, whereas zapodids show none. The for as long as 6 months in Wisconsin, is lower jaw is remarkably different in the two another interesting trait. groups. The characters in common, admit- In Wisconsin, the grooved incisors alone tedly important, are the enlarged antorbital would identify the two zapodid species, ex- foramen and the peculiar lacrimal-jugal ar- cept that Reithrodontomys and Synapto- ticulation (Lyon, 1901). But the foot struc- mys also have grooves. No mice are likely ture alone, with a tri-radiate “cannon bone” to be confused with the long-tailed, long- of the hind limb specialized far beyond that legged zapodids. of any Artiodactyla, would separate Zapus None of the other Wisconsin mice shows from Dipus as pigs from camels, so it would such yellowish-ochraceous tones in the dor- seem. As Lyon mentioned early on, and sal pelage. The much folded enamel surfac- many recent workers have re-affirmed, the es of the molar teeth are also characteristic, Zapodidae show affinity to the Muridae (My- the labial border of each elaborated and de- omorpha) regardless of their hystricomor- fined by re-entrant “bays” (see Fig.) as sev- phous antorbital canal. eral arcuate cusps and crests. There are four The Zapodidae range across two con- along the border of MI or of M2 (these cusps tinents, from Europe eastward through Si- are less arcuate in Napaeozapus, especially beria (subfamily Sicistinae, birch mice), the second cusp for each tooth, which is not southward in China (Zapodinae, genus so isolated but is confluent with the mid-por- Eozapus), and across North America in suit- tion of the tooth surface). There is one lin- able habitats of Canada and much of the gual (inner) re-entrant groove on each of United States (Zapodinae, Zapus and Na- these molars. Also of interest is the jugal, paeozapus). The American jumping mice expanded and elevated anteriorly so as to have elongated tail and hind legs and are join the maxillary at a vertical suture, and able to leap incredible distances (in Napae- even to contact the lacrimal bone in the or- ozapus as far as two meters). Unlike some bit. The auditory bullae are vase-like, each other leaping or saltatorial mice, the audi- opening by a large meatus. The two premax- tory bullae are not enlarged, nor are the illaries fuse between the upper incisors and ankle bones fused. There are no external form a keel, and the upper incisors are or- cheek pouches, and the molar teeth are ange anteriorly but the orange overlies ivo- rather hypsodont (unless worn smooth). ry-yellow enamel behind the surface.

304 THE WILD MAMMALS OF WISCONSIN Key to Wisconsin Jumping mice Zapus hudsonius (Zimmermann) Meadow Jumping Mouse 1 Tail almost always lacking a white tip, color of fur on sides yellowish tan, up- 1780. Dipus hudsonius Zimmermann. Geog. Ge- per premolar present and of reduced size, schichte die Mensche vierfussigen Thiere. 2: labial cusps of upper molars distinctly 358. Type locality Hudson Bay. arcuate, the second on M-l/ and M2/ 1781. Dipus labradorius Kerr. Animal Kingdom. isolated by re-entrant groove, tail usually Page 276. Based on description by Pennant. scaly, caudal hairs not conspicuous ..... 1822. Gerbillus canadensis Desmarest. Mammalo- ...... Meadow Jumping Mouse gie, 2: 321. Zapus hudsonius 1825. Gerbillus labradorius Harlan. Fauna Ameri- 1’ Tail showing a more or less distinctive cana. Page 157. hirsute white-tipped end, color of dor- 1799. Dipus americanus:Barton. Trans. Amer. sum and especially of sides bright but Philosolphical Soc.,4: 115. darker tawny ochraceous orange, upper 1829. Meriones labradorius: Richardson. Fauna premolar lacking, labial cusps of upper Boreali-Americana, 1:144. molars indistinct, the second on M1/or 1843. Jaculus labradorius Wagner. Suppl. M2/ confluent with mid-surface of tooth, Schreber’s Saugthiere, 3: 294. tail scales intermixed with conspicuous 1899. Zapus hudsonius hudsonius: Preble. N. Amer. hairs, especially beyond hind feet ...... Fauna, 15: 15...... Woodland Jumping Mouse Napaeozapus insignis The name Zapus means, “Oh, what a big foot” referring of course to the hind foot. The trivial part of the binomen, hudsonius, Genus Zapus Coues refers to Hudson Bay, although no type spec- Jumping Mice imen exists from Hudson Bay. Anderson (1942: 37) chose Fort Severn, Ontario, as a “He glides around the fire much as the others do, locality from which typical jumping mice can but at the approach of danger, he simply fires be obtained. himself out of a catapult, afar into the night. Eight Description. The characteristics were set or ten feet he can cover in one of these bounds and forth above for the family Zapodidae. In com- he can, and does, repeat them as often as neces- parison to Napaeozapus, the meadow jump- sary... Since the creature is chiefly nocturnal, the ing mouse averages smaller and the baculum traveller is not likely to see it, excepting late at night is shorter. The tip is flattened (Burt, 1960). when venturesome individuals often come creeping Upper premolars are present. The number about the campfire... what is the reason for this indeterminable tail? The answer is, it is the tail to the kite, the feathering to the arrow; and observation shows that a Jumping Mouse that has lost its tail is almost helpless... one individual de-tailed by a mowing machine had no control of the direction, and just as often as not went straight up or landed wrong end to, and sometimes back where it had started from.” — Ernest Thompson Seton, in Ghosts of the Campfire, 1913.

Sketch of Zapus hudsonius by Wm. E. Hitchcock.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 305 of chromosomes is 2N =72, which is a rather found them more abundant in Minnesota on large number (Hoffmeister, 1989). There are mima-like mounds than on surrounding wet- four pairs of mammae. lands, probably as these higher sites are more The dorsum is furred with guard hairs and suitable for burrows. underfur, a mixture of ochraceous or yellow- The meadow jumping mouse makes a nest ish and dark brown or black. The longitudinal of dry grasses and leaves about 6-7 inches in band is darkened by the black hairs, which are diameter with one entrance on a side (Dexter, fewer and less visible on the sides, and lacking 1954; Jackson, 1961). It may be placed in altogether in the lovely lateral band (not always dense vegetation or shallow excavations. In present) of golden ochraceous or yellow adja- winter the hibernation nest is situated in deep- cent to the white underparts. The tail is bicol- er tunnels (two feet or even deeper). The hi- or, blackish above (often grayish or brownish), bernation nest may contain a few leaves. pale whitish below. The large eyes are black. Foods. Little is known about the foods The upper surfaces of the feet are white or of Zapus in Wisconsin. Whitaker (1972) sum- grayish white. Rarely the mice show white spot- marizes several studies including his own. ting, and one was melanistic. Occasionally they Hamilton (1935) mentioned berries and nuts have white-tipped tails (Whitaker, 1972; as well as seeds, fruits, and insects. Whitaker Schorger, 1951). The upper incisors are or- (1963) reported on 796 individuals from New ange or yellow. York, finding that insects make up about half Although most workers conclude the the food in spring, whereas seeds are only sexes are the same size, Whitaker (1972) 20%. As summer passes seeds become more and Whitaker and Wrigley (1972) report the important, animal foods less, and the fungus females are slightly larger. The total length Endogone becomes an important compo- is about 200 to 256 mm, and the tail ap- nent. The most important animal foods then proximately 60 percent of total length. See were lepidopterous larvae and carabid and Table Rod-15. curculionid beetles. In Indiana, in a sample of Dental Formula. I 1/1, C 0/0, P 1/0, M 131 individuals, Whitaker (1970) found that 3/3 =18. A reduced P4 is present, and the seeds comprise most of the diet (50.4%), the upper incisors are distinctly grooved. See Fig. species depending on the season. Impatiens under Mus. and Setaria were important by volume and Geographic Range and Status. This ben- frequency. Lepidopterous larvae, insects and eficial, beautiful mouse is state-wide (see Map). slugs were also eaten. Habitats. Often found in wet situations, Reproduction. Quimby (1951) found the such as swamp and marshes, the meadow gestation period to range between 17 and 21 jumping mouse will occasionally be found in days for four females. He concluded that in well-drained meadows and mature forest. The Minnesota most births take place between 15 meadow jumping mouse is abundant in grassy June and 30 August, but peaks occur in late shorelines of streams and ponds. Whitaker June, mid-or late-July, and in mid-August. Preg- (1963) mentions the importance of herbaceous nancies are noted by several workers in every ground cover, and Getz (1961) reported that month of summer and early fall, but there is the mouse avoids sparse vegetation. Other more breeding in June. The latest litters were reports on ecology are found in Quimby (1951) September. Most females produce two litters. and Hamilton (1943). In Wisconsin I have of- Quimby (1951) found an average of 5.7 young ten taken Zapus in marsh grasses, jewel weed, per litter (N = 17, range 4-7) in Minnesota, and dense grass near tamarack swamps, and and Townsend (1935) reported 4.5 (N=17, 3- have observed them occasionally in open for- 7) for New York. Krutzsch (1954) reported a est, on both wet and dry soils. Ross et al. (1968) mean of 4.5 (range = 2-8) from over a wide

306 THE WILD MAMMALS OF WISCONSIN Upper molar row of Zapus. After P. Krutzsch, 1954.

Maps showing geographic distribution of Zapus hudsonius in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 307 geographic range. Whitaker (1963) reported white. By three weeks the body was haired 5.5 (N = 78) with a range of 2-9. out. The eyes had opened in the fourth week. In Wisconsin specimens, few pregnancies Adult pelage had appeared approximately at were noted. Lactating or pregnant females this time, replacing the juvenal fur. The inci- were obtained from June into July in Portage sors had become orange, and the tail had County, and ranged from June into October grown 10-fold. Growth slowed thereafter, and judging from other records in Wisconsin (June adult size was attained in 60 days or more. 23 in St. Croix County; through July, in Ad- Mortality. Many predators prey on jump- ams County; July 9 in Ashland County; 25 ing mice, including frogs, snakes, fishes, fox- September near Pittsville, Wood County; and es, wolves, weasels, mink, and house cats. 27 October near Necedah, Juneau County). Birds of prey include several hawks, especial- Breeding seemed to last through September ly red-tailed and marsh hawks, as well as sev- but occurred only sporadically after July. None eral kinds of owls (Whitaker, 1963; Baker, was observed in August in our specimens. Nora 1983). Known parasites are well summarized Lopez-Rivera found three records of pregnan- by Whitaker (1972), and reportedly mites are cy from the Buena Vista Marsh in the Hamer- the most common external parasites. Several strom’s field journals. They were June 2, July species of fleas, botflies, chiggers, sucking lice, 26, and August 2. and mites were also reported. Internal para- Thus, the pregnancies for Zapus in Wis- sites included trematodes, tapeworms, and consin occur in every month, from early June nematodes. Carmichael (1979) reported blood until late October (1 record). Litter size aver- flukes, Schistosomatium douthitti, from aged 6 (range 5-8) for six pregnant females. I jumping mice in Minnesota. reared one live litter of five wonderful young Home Range and Density. Quimby [on milk-egg food] in Portage County. On the (1951) reported home range from 0.19-0.87 Buena Vista Marsh, 3 females contained 6-7 acres for females, and 1.1-0.14 acres for embryos (Hamerstrom records). Altogether males, in Minnesota. In another study he the 10 females averaged 6.2 (5-8) embryos. found the males to have larger home ranges Juvenal mice entered the population in July, (2.7, 9 males and 1.57, 17 females). In Mich- and again in early fall (specimens taken 20 igan, Blair (1940) determined the home range September in Dodge County; 19 September as about the same for males and females (0.89 in Columbia County; 22 October in Vernon to 0.92 acres, for 26 males and 24 females). County; and 10 October in Oconto County). Quimby (1951) and Muchlinsky (1988) Jumping mice were taken as late in the fall as suggested that for the most part the popula- September 22, 11 October, 18 October, and tion turned over in summer, the new adults 27 October, after which the mice enter hi- replacing the carry-overs from the year previ- bernation. ous. The populations vary in size from year Whitaker (1963) discussed ontogeny of to year, but two studies agree fairly well, plac- 19 newborn young. The young measured 34.4 ing the number of meadow jumping mice at (range 30-39) mm in total length, the tail 9.3 about 12 per acre (Quimby, 1951; Blair, (7-11), hind foot 4.7 (3-6). The average weight 1940). The habitats for these mice are not for 14 neonates was 0.8 g (0.7-1.0 g). Quim- far removed from Wisconsin. by (1951) described the newborn young as pink Remarks on Hibernation and Behavior. and naked with minute vibrissae. After about The solitary meadow jumping mouse is usu- a week the vibrissae became visible to the na- ally nocturnal, but often seen by day. It can ked eye, the tail became bicolored, the pinnae climb, and occasionally does to cut seed heads of the ears unfolded, and claws appeared. The off tall plants such as timothy (Whitaker, per- incisors erupted on the 13th day. They were sonal correspondence). It is an able swimmer,

308 THE WILD MAMMALS OF WISCONSIN Table Rod-15. Cranial and external measurements and weights (g) of Zapus hudsonius.

Greatest Condylo- Zygomatic Inter Length Maxillary Length Total Length Length Length Weight length basal breadth orbital nasals tooth Incisive Length Tail Hind ear grams skull length breadth row foramen Vertebrae Foot intermedius St. Croix Co. 23.03 20.86 10.86 4.22 8.30 3.72 4.42 202.25 121.50 28.25 9.75 ±0.66 0.21 0.52 0.22 0.87 0.33 0.29 14.13 ±8.89 2.22 1.71 N=5.00 5.00 5.00 5.00 5.00 5.00 5.00 4.00 N=4.00 4.00 4.00

Sauk Co. 22.05 20.10 10.27 4.06 8.54 3.76 3.91 196.30 121.00 27.30 10.30 ±0.90 0.97 0.18 0.17 0.54 0.22 0.32 N=7.00 7.00 7.00 7.00 7.00 7.00 7.00 3.00 3.00 3.00 3.00 hudsonius Bayfield Co. 22.37 20.80 10.64 4.21 8.45 3.89 4.20 205.82 124.94 29.29 9.88 17.83 ±0.65 0.67 0.34 0.16 0.58 0.19 0.22 6.47 ±6.47 1.45 0.96 3.00 N=20.00 20.00 20.00 20.00 20.00 20.00 20.00 20.00 N=20.00 20.00 20.00 17.00

Portage Co. 22.05 20.01 10.53 4.13 8.24 3.79 4.24 202.40 121.58 29.34 12.44 20.50 ±0.75 0.87 0.31 0.16 0.42 0.22 0.21 2.87 ±4.86 0.91 1.38 N=16.00 16.00 16.00 16.00 16.00 16.00 16.00 13.00 N=13.00 13.00 13.00 13.00 for several minutes at least, swimming frog Muchlinsky (1988) measured the soil tem- style either with head out of the water, or with perature at 50 cm depth to estimate emergence head and body completely submerged. It can temperature for Zapus hudsonius during the leap usually about 1 meter for the initial jump, years 1977-1988. Males emerged slightly ear- which helps them escape predators. Once I lier than females (males April 20-May 11, and attempted to leap on one, and it was 20 feet females May 6-7, until May 29). The soil tem- away before I landed (my knee upon a sharp perature varied from 8-15 degrees C. rock). The jumps are much shorter after the Additional Natural History. Whitaker initial leap. Two-meter leaps have been re- (1972) reviewed the biology of Zapus hud- ported as errors, but Jackson measured a leap sonius. slightly greater than that. Geographic Variation. There are two Hibernation of Zapus is interesting, geographic races recognized in Wisconsin which in Wisconsin is genuine and seems to (Krutzsch,1954). He drew a line across the last from late October until mid-April. The northern parts of Wisconsin assigning all spec- physiological data on hibernation have been imens north of it to the nominate race, which well summarized by Whitaker (1972). A litter mice are darker especially on the sides. Jack- I reared surprised me, because the young mice son (1961) drew the line somewhat different- ate the eyes of one that had begun to hiber- ly, probably intuitively and hopefully ascrib- nate, and ate another entirely except for the ing races to the lake country in the north and skin. During the summer the young were the prairie country in the southwest and south- amiable to one another, and to me, but as east (near Green Bay). He changed one mar- hibernation approaches in September, they ginal record from Oconto County from the must be separated. southern race to the northern. Inasmuch as

TAXONOMIC ACCOUNTS / ORDER RODENTIA 309 the cranial measurements do not vary greatly over Wisconsin, and the traits of skull and ly, nor seem to be diagnostic in Wisconsin, I color vary in a spurious microgeographic attempted to assign the races on the basis of and confusing manner. Concerning cranial color, the tawny ochraceous ones southward characters, Krutzsch (1954) lists for Z. h. and the dull-sided darker ones northward. I hudsonius the auditory bullae broader (less only compared specimens of large size be- inflated), condylobasal and zygoma longer cause young animals are never as bright as on average, braincase broader, and incisive the older ones. The results (see Fig) show some foramina shorter even though the overall variation in color, as would be expected, but size is larger. In Wisconsin specimens of in- fair constancy in the northern and central termedius approaching hudsonius, he not- populations. It is necessary to shift the bounded longer bullae and wider bowed forami- ary southward in the central counties, for na. In these characters there is much overpopulations there are decidedly dark. lap in Wisconsin. A series of intermedius Krutzsch (1954) made his assignment from St. Croix County has small skulls (Ta- based on a huge taxonomic revision, but in ble Rod-15) and the incisive foramina are Wisconsin the specimens were rather few. narrow (less bowed). The Sauk County in- Jackson made a sensible assignment owing termedius are similar but have relatively nar- to his knowledge of the ecology of Wiscon- row foramina. The large Drummond series sin, but my specimens in the central counties (Bayfield County) shows larger dimensions (Marathon, Portage, Juneau) do not fit either and wide-bowed foramina, resembling the arrangement. I provisionally redrew the Portage County series. Juneau County skulls boundary line between the races (see Map). resemble hudsonius, whereas Adams Coun- In cranial characters I could not distinguish ty skulls east of the Wisconsin River resem- significant differences on which to draw sub- ble intermedius. species boundaries, and color also had limited use. Inasmuch as color in Z. h. hudsonius is reportedly darker and more tawny on the sides Zapus hudsonius hudsonius (Zimmermann) (i.e., more light brownish orange instead of yellowish) than in intermedius, I composed two 1780. Dipus hudsonius Zimmermann. Geogra- series of comparative skins including no. 3775 phische Geshichte... 2: 358. Type from “Hud- Adams Co. and 5835 Marathon County for son Bay” but now fixed at Fort Severn (see text). orange (the latter darker) and for black inter- 1852. Meriones americanus: Lapham. p.44. Type mixtures on the sides 7082 Pierce Co., 4650 from near Philadelphia, as Dipus americanus Adams Co., 5837 Bayfield Co., 5839 Bay- Barton, 1799. field Co., the last darkest. These were weight- 1899. Zapus hudsonius hudsonius: Preble. N. Amer. ed 1 to 2 and 1 to 4, respectively, and the Fauna, 15:15. samples were averaged with the two values added (large values darkest). On the whole, the Description. Pelage dark, sides dull and earlier observation that specimens are dark in less tawny, auditory bullae larger, incisive fo- the central counties as well as in the north was ramina broad. borne out. In Marathon and Portage counties Geographic Range. This race is found in and some localities westward, the dark color is northern counties. See Map. due more to the presence of orange, and in Specimens examined. Total, 109. Ash- Bayfield County and other localities black hairs land, Bayfield (includes Outer Island, Apos- cause dark color. tle Islands 1), Douglas, Florence, Iron, Ju- The race Z. h. intermedius is not al- neau, Langlade, Lincoln, Marathon, Ocon- ways distinctive, intergradation occurs wide- to, Oneida, Portage, Wood counties.

310 THE WILD MAMMALS OF WISCONSIN Zapus hudsonius intermedius Krutzsch of curious form and habits, hibernates in winter, and is exceptionally beautiful. 1954. Zapus hudsonius intermedius Krutzsch. Univ. Specimens examined, Total , 72. Adams, Kansas Publs., Mus. Nat. Hist., 7(4): 447. Type Buffalo, Clark, Columbia, Dane, Dodge, from Ridgeway, Iowa. Iowa, Manitowoc, Monroe, Pierce, Sauk, Sheboygan, St. Croix, Vernon counties. Description. Pelage brighter tawny on Additional Record. Marlin Johnson average especially on the sides. Measurements caught a meadow jumping mouse in extreme are given in Table Rod-15. northern Door County. Geographic Range. This race of jumping mouse occurs in suitable habitats though- out much of southern Wisconsin. See Map. Genus Napaeozapus Preble Status. This mouse of marsh, swamp, Woodland Jumping Mouse meadow and forest avoids cultivated fields, and has never been considered a pest. It can The genus Napaeozapus differs in several be considered beneficial to humankind, be- characters from other zapodids and is thought cause it feeds on plant seeds, many consid- to be evolutionarily more advanced. Resemb- ered as “weeds”, and on insects, many of es Zapus hudsonius, especially in the south- which are pests. It is a prey species of numer- ern parts of its range (where bright colored ous carnivores and birds of prey. The num- Zapus are common), the woodland jumping bers of specimens indicate the mouse is un- mouse tends to be more brightly orange or common, but it can be locally abundant. This ochraceous on the sides and in the pure ochra- mouse also has much aesthetic value, for it is ceous lateral line (when present). It has the terminal portion of the tail hirsute and white (5-20 mm). The hairy tail is less scaly in appearance and more sharply bicolor; the overall size is larger on average; there are no premolars present; and the upper Ml and M2 show less conspicuously arcuate labial cusps (i.e., less prominent re-entrant grooves defin- ing the cusps. There are 72 chromosomes as in Zapus (Whitaker and Wrigley, 1972).

Napaeozapus insignis (Miller) Woodland Jumping Mouse

1891. Zapus insignis Miller. Ame. Nat., 25: 742. Type from Restigouche River, New Brunswick. 1899. Napaeozapus insignis: Miller. Bull. New York State Mus. Nat. Hist. 6: 330. The generic name refers to the animal’s Color scale values for Zapus hudsonius in Wisconsin. habitat: woodlands. It also exclaims, “Oh, Under each value is the sample size. Dark pelages are mostly what a foot,” referring to the hind foot. The in the northwest and center. This finding roughly agrees with Krutsch 1954, but the variation is irregular and the specific name insignis means conspicuous, differences not distinct. as in insignia, and doubtless refers to the

TAXONOMIC ACCOUNTS / ORDER RODENTIA 311 white-tipped tail. The description is given Napaeozapus. Whitaker and Wrigley (1972) above for the genus. conclude that Napaeozapus inhabits the spruce-fir and hemlock-hardwood forests in the eastern half of North America, especially Napaeozapus insignis frutectanus Jackson balsam fir and eastern hemlock. Locally, the mouse prefers cool, moist habitats within the 1919. Napaeozapus insignis frutectanus Jackson. forest or at its edge. It also occurs, as noted Proc. Biol. Soc. Washington, 32: 9, February above, in bogs and swamps, but occasionally 14, holotype from Crescent Lake, Oneida inhabits dryer areas (Sheldon, 1934). Damp County, Wisconsin. rocky areas are often preferred. Preble (1956) noted that the largest populations in his expe- Description. See above. Napaeozapus rience were found along streams in grasses, averages slightly larger than Zapus hudso- sedges, and alders. Whitaker (1963) noted a nius. See tables Rod-15-16. preference for low herbaceous vegetation. Dental Formula. I 1/1, C 0/0, P 0/0, M Although often taken together, Zapus and 3/3 = 16. Napaeozapus may segregate in different plant Geographic Range. The woodland jump- communities, with Zapus primarily in mead- ing mouse is found in the boreal or northern ows and Napaeozapus primarily in woods. parts of Wisconsin (see Map). Frank Iwen iden- Hamilton (1935), Preble (1956), and others tified a specimen (UW 24342) from Fort Mc- found Zapus invading Napaeozapus habitats, Coy, southwest Wisconsin. I examined the but not the other way around. My trapping in specimen; its tail was not tipped white, but Wisconsin has yielded results consistent with the color was rich orange brown. Anterior that view. Little is known about dens, nests, premolars were reportedly absent; we could and burrows of Napaeozapus. Apparently the not find the skull. mouse usually places its nest in a burrow, but Status. This jumping mouse is and always occasionally makes it in low vegetation. Stup- was rare, known in Wisconsin only by 14 spec- ka (1934) found a nest made of leaves and dry imens in 1961 (see Jackson, 1961). The num- grasses in a brushpile in Maine, and Snyder ber of specimens has about doubled, but the (1924) describes a natal burrow excavated near species remains rare in comparison to most Lake Nipigon, Ontario. The burrow extended Wisconsin mammals. Napaeozapus is not a about 5 1/2 inches below the soil, for a length pest, and owing to its beautiful colors and clean of about 1 meter, before ending in a blind graceful appearance its habitats should be pre- pocket. The nest cavity was about 14 inches served. It is an interesting mammal, because it (= 355 mm) from the entrance, about 6 by 5 hibernates and is evolutionarily adapted for inches, and contained a nest of dry leaves. jumping. Napaeozapus is more colorful and a There were five young in the nest. more spectacular leaper than Zapus. Foods. Nothing is known of food habits Habitats. Occurs in northern forests, in- of Napaeozapus for Wisconsin, but they are cluding maple, pine, spruce, tamarack, cedar, probably similar to those elsewhere, and, and similarly wet and boreal woodlands. John where the two species coexist, probably sim- Long and I have taken them at the edge of ilar to food habits of Zapus. Before the fun- cedar-tamarack swamps near standing water, gus Endogone was known to be a food for rusty with mineral deposits, and in jewel weed, jumping mice, the foods were classed as most- mosses and ferns on black wet soil away from ly seeds and insects. Most (some 70%) of the the standing water. My students have taken foods were seeds, depending on their avail- them near small streams in northern Wiscon- ability in season, and berries. About 30% of sin. Zapus is often taken in association with the diet consisted of insects and soft-bodied

312 THE WILD MAMMALS OF WISCONSIN Upper molar rows of Zapus and Napaeozapus. After Krutzsch.

Maps showing geographic distribution of Napeozapus insignis in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 313 invertebrates (Hamilton, 1935; Sheldon, and parturition continues sporadically through 1934; Whitaker, 1963; Wrigley, 1972). Lar- July. A second peak was apparent in August val Lepidoptera, adult beetles, larval flies, spi- (in sharp contrast to our Wisconsin data for ders, centipedes, worms, and other insects Zapus) and many of the females produce sec- are eaten. Some leaves and shoots are con- ond litters. Late season pregnancies were in sumed (Whitaker, 1972). Endogone may August, and the latest observed September make up 35 percent of the diet (Whitaker, 1. In second litters, litter size is reduced. Sec- personal correspondence). ond litters are less often produced in the north- Reproduction. The number of woodland ern latitudes (Wrigley, 1972). As in Zapus, a jumping mice taken recently in Wisconsin al- turnover of the carry-over adults occurs. The most doubles the 14 specimens (from 10 lo- old mice are replaced by young of the year calities) discussed by Jackson (1961). Nev- (Muchlinsky, 1988). ertheless, breeding information is meager. Newborn young are hairless and pink with One lactating female (2207) from Oconto loose skin (Lane and Hamilton, 1954), white- County was obtained July 22, 1970, and a tipped tail prominent (Snyder, 1924). Vibris- young Napaeozapus (sex?) was obtained 7 sae are visible and the tiny ear pinnae folded. August 1971, from Chippewa Lake in Bay- Measurements reported were: total length 35.2 field County. Most of the individuals exam- mm, tail length, 11.0, hind foot length 5.0. ined were males. After a week the vibrissae were 2 mm long From outside Wisconsin the gestation and the ear pinnae larger. By 10 days, pig- period was roughly determined as about 29 ment spots were visible and the ear pinnae had days or even longer. Quimby (1951) suggest- unfolded. By 14 days fine hair covered the ed a shorter time. Whitaker and Wrigley body. Lower incisors were visible by 19 days. (1972) suggest 23-25 days, based on lab By 21 days the dark dorsal band on the fur matings. Other gestation periods estimated appeared. Eyes opened by 26 days. The inci- were 23 and 25 days. Napaeozapus seem- sors were grooved and yellow by 31 days. By ingly are not sexually active until their second 34 days, the young resembled adults except year. Eighty litters from a wide geographic the pelage was paler, more yellowish rather range averaged 4.6 (2-7), and 45 counts of than orange-brown. The adult pelage appeared placental scars averaged 4.3 (2-7). Apparently between 63 and 80 days (Wrigley, 1972). Napaeozapus has no postpartum estrus, al- Mortality. Predation on woodland jump- though late breeding may occur after the ing mice by carnivores, snakes, and birds of young are weaned (Schwentker, 1939). prey is important. Fish may swallow them if Pregnant mice were collected in early they fall into a pond or stream. Some records May, although only a few. Births peak in June of predation include screech owl, mink, wea-

Table Rod-16. Cranial measurements of Napaeozapus.

Greatest Condylo- Zygomatic Inter Length Maxillary Length Total Length Length Length Weight length basal breadth orbital nasals tooth Incisive Length Tail Hind ear grams skull length breadth row foramen Vertebrae Foot Wisconsin 24.26 21.83 12.17 4.62 9.50 4.02 4.89 230.17 142.27 30.42 13.33 20.95 & ±0.42 ±0.51 ±0.25 ±0.42 ±0.42 ±0.17 ±0.2 ±6.64 ±3.07 ±0.9 ±2.5 2.27 N=7 N=7 N=7 N=7 N=7 N=7 N=7 N=12 N=12 N=12 N=12 N=2

Cheboygan Co., Mich. 24.0 21.3 11.7 4.45 8.5 4.4 5.0 233 142 31 16

314 THE WILD MAMMALS OF WISCONSIN sels, striped skunk, domestic cat, bobcat, and Co.: Fort McCoy 44.01N, 90.4W 1 UW (skull timber wolf. Internal parasites include proto- not seen). Portage Co.: 1 mi. E Jordan Pond, zoans, tapeworms, and roundworms. Exter- in Jordan Swamp (1/4 mi. N Hwy 66) 1. Price nal parasites include fleas, mites, ticks, and Co.: T32N, R2W, Sect. 9, 1 Hamerstroms. botflies (Wrigley, 1972). Sawyer Co.: 1/2 mi. N Couderay 3. Taylor Home Range and Density. Home range Co.: 16 mi. S Medford 1. Wood Co.: Mead seems about 1.0 to 6.5 acres in females, and in Wildlife Area 1. males 1.0 to 9.0 acres according to Blair (1941). Michigan. Cheboygan, 5. In New York, Napaeozapus are fairly abundant; Townsend (1935) found as many as 24 per acre. In Michigan Blair found 2.7 per acre, and Man- Family ERETHIZONTIDAE Thomas ville (1949) estimated 0.26 per acre. Brower New World Porcupines and Cade (1966) estimated 5.2 per acre in New York. Whitaker (1963) observed frequencies up Form stout, forearms robust, dorsum with long to 0.21 per 100 trap nights. guard hairs, wooly under fur, and conspicu- Remarks on Behavior and Hibernation. ous sharp quills; molars flat-crowned, rooted, Usually after several leaps the woodland jump- and hyposodont; angular process of dentary ing mouse hides in vegetation unless closely inflected; infraorbital canal larger in diameter pursued. It climbs in bushes, but has never than foramen magnum of spinal cord; planti- been seen to ascend a tree. They swim with grade feet specialized for a clambering, arbo- head above or below the surface (Wrigley, real niche. Nevertheless, porcupines are of- 1972). Usually nocturnal, they have been seen ten on the ground. The origin of this hystri- in daylight. Late records, preceding hiberna- comorph family was in Central or South tion, are in late September for adults, and for America, but it, like the opossum, ranged young-of-the-year October. Wisconsin speci- northward, and today porcupines are on the mens show collection dates from 10 May until Arctic slope of Alaska. late September, with one adult taken 16 No- vember. These mice put on weight before entering their hibernaculum, and subsist on fat until they emerge in early May or, in females, a little later. The earliest emergence date known is 16 April in Pennsylvania. Geographic Variation. Minor variation was seen in the length of the white on the tail, which in Portage and Marathon counties seemed reduced, and in one specimen from Pulaski Road, Marathon County, lacking. There is one race in Wisconsin. Specimens examined. Total, 24. Bayfield Co.: Clam Lake Field Station 2. Chippewa Lake Field Station 4. 3 mi. NW Clam Lake Field Station 1. 1 mi. N Grandview on 10- mile Creek 1. Drummond 1. Douglas Co.: St. Croix River 2. Forest Co.: Laona 1. Langlade Co.: Summit Lake, Camp Susan 1. Marathon Co.: 1/2 mi. N on Pulaski Road 2. Marinette Co.: 9 mi. NW Crivitz on Co. A, 1. Monroe Sketch of Porcupine in tree. By Walter A. Weber.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 315 Genus Erethizon F. Cuvier In winter the hair on the shoulders grows out North American Porcupine longer. The quills are spinose, with black tips. The limbs are stout, as are the claws, and the Races reviewed by Anderson and Rand, Canadian J. plantigrade feet have exceptionally broad Res., Sec. D, 21:292-309, 1943. soles, naked and scaly. There are five toes on 1822. E [rethizon]. F. Cuvier. Mem. Mus. Hist. Nat. the hind foot, four on the forefoot. The belly Paris, 9:432. Type Hystrix dorsata Linnae- is covered with sparse blackish or brown hair us, 1758, Syst. Naturae. (no quills). The face is furry, and the eyes fairly large. The ears are small and the rhinarium Description. See account of Erethizon- broad. The tail is bristly on the lower sides. tidae above. The Erethizon have a well-devel- There is a distinctive but variably formed bac- oped hallux, but the tail is not prehensile. The ulum, with a club-like base and flattened (dor- related South American Coendu is the oppo- soventrally) shaft (see Burt, 1960). site. Normally a single young is born per year. Jackson (1961) reported six teats, four Young are born with hair and quills enclosed pectoral and two abdominal, but Burt (1948) in the amnion. The genus is monotypic. and Baker (1983) report only four pectoral mammae present. Uldis Roze told me that perineal glands, paired pockets near the ven- Erethizon dorsatum (Linnaeus) tral midline, larger in males, are likely used in dragging and leaving odors (produced by se- 1758. [Hystrix] dorsata Linnaeus. Syst. naturae, 10th baceous secretions and bacteria (Staphylococ- ed. 1:57. Type locality Eastern Canada. cus, Cryptococcus)). 1822. E[rethizon] dorsatum: F. Cuvier. Mem. Mus. The porcupine skull is large for Wiscon- Hist. Nat., Paris, 9:432. sin mammals, with formidable orange incisors. The high, flat-crowned cheek teeth show a The word Erethizon means something peculiar folded molar pattern. The nasals do of an irritation, and the word dorsatum re- not extend as far forward as the incisors, al- fers to the upper parts or back, where the though the fleshy nosepad does. The interor- formidable quills are found. bital breadth is quite wide. It is over 50 percent of the greatest breadth of the zygomata, which zygomata narrow anteriorly. The in- Erethizon dorsatum dorsatum (Linnaeus) fraorbital canal is enormous, larger even than the foramen magnum. The angular process Description. See remarks above under Erethizontidae. The porcupine is a medium- sized mammal, but is an exceptionally large rodent. Among Wisconsin rodents, only the beaver Castor exceeds it in size. The neck is so short it seems absent, and indeed the anterior two cervical vertebrae are compressed and fused together. The tail is short and well armed with quills around the border, which quills are slapped in the face of an enemy. The shaggy whitish-yellow guard hairs and long yellowish quills give the porcupine an appearance of prickly, yellow and black, plumpness. The longest quills exceed 60 mm. Skull of Erethizon dorsatum.

316 THE WILD MAMMALS OF WISCONSIN of the dentary is inflected. The sagittal ridges converge near the occiput to create a sagittal crest in adults. The palate does not extend posteriorly as far as the last molars. The ground color of the porcupine dorsally is black or dark brown, with stout yellowish or ivory hollow quills intermixed with grayish or whitish guard hairs and dark wool. In winter the shoulder hair greatly lengthen. The underside is dark brown. Jackson (1961) reported albinos from Wisconsin. Two from Wisconsin were placed in my freezer lacking, unfortunately, any additional data. Molt takes

Maps showing geographic distribution of Erethizon dorsatum in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER RODENTIA 317 place in late spring and summer (Po-Chedley Jackson’s (1961) line agrees with mine and Shadle 1955). (for the present distribution). I mentioned The porcupine varies to 900 mm (35.5 in) above that there are no porcupines on islands in length, weight to 30 lbs (13.5 kg). The skulls in Door County, but porcupines still exist in average 100 mm (approx. 4 in) in total length. fair numbers on the forested Peninsula north Dental Formula. DF = I 1/1, C 0/0, P of Sturgeon Bay. Porcupines regularly occur 1/1, M 3/3 = 20. in northern Portage and Wood counties. They The 20 teeth include four upper molari- seem absent from southern Portage County. form teeth. The dental formula distinguishes the Porcupines possibly occur in Monroe and porcupine skull from the flat-skulled marmot, Juneau counties on occasion, and two were which has prominent postorbital processes. dead on Highway 10 at Wedgee and Five- Geographic Distribution. This boreal mile creeks, in Clark County. See Map. mammal ranges or did range northward as far Status. A century ago many American as the Arctic Slope of Alaska, and southward children were taught to “Save the porcupine” along the Rockies into Mexico, and southeast- because a lost person in the North Woods ward into west Texas. All of Michigan, Wis- can always capture one and eat it. That is a consin, Minnesota, most of Iowa and Indiana, fanciful legend. Perhaps porcupines are as and northern Illinois lie within the probable rare as people who get lost in the North geographic range (Hall and Kelson 1959). Woods, at least in the remote forests wherev- Apart from archaeological records (Parmalee er they happen to get lost. The porcupine is 1967, 1963), there is only one historic record edible (Jackson 1961). It is not uncommon in of porcupines in northern Illinois. Kennicott Upper Michigan. Many people consider the (1859:91) mentioned a record from Whiteside porcupine a nuisance. It damages trees by County and the banks of the Illinois River. feeding on the leaves, bark, and buds, and Neither he nor Hoffmeister (1989) accepted sometimes kills trees. It gnaws on cottage this record. Lyon (1936) listed records from decks, eats plywood, chews on cabin walls, Indiana near the Illinois border, and Bowles and even rubber tires, and occasionally cuts (1975) thought the species was probably in electric wires. It is active day or night. The northeastern Iowa. There are historic reports observed damage seems important, but in the from southern Wisconsin (Jackson 1961). Now big picture it is insignificant. Injury from quills there are no populations remaining there. may inflict great pain and injury to dogs, nat- Deforestation and land use have devas- ural predators, and even humans. Quills tend tated porcupine numbers in the former hard- to work inward, not outward, and I personal- wood forests south of the “tension zone.” The ly know of rare mortality to black bear and northern forests still support this mammal in elk. Doubtless some coyotes and dogs die from fair numbers. Jackson (1961) noted the re- quill injuries on occasion. Quills may be retreat of the species, listing authentic records moved with pliers, but the quill must be loos- “before 1870” in Richland and southern Sauk ened by gentle side-to-side movements, close counties. He mapped reports from Walworth to the wound, with great caution used to pre- (1917), Sheboygan (1917), and Dane coun- vent breaking the quill. Snowshoe hares and ties, and westward (Buffalo County) along the deer eat porcupine “leavings” (i.e., dropped Mississippi. He examined a specimen from plant parts) dropped from a tree in winter. Monroe County, and proposed 1961 margin- Bears, fishers, and other carnivores occasional limits as extending from Burnett, into Rusk, ally kill and eat their ventral parts. The fisher is Clark, and Shawano counties, and northern considered the primary controlling predator of Door County. No records are known now from porcupines. Native Americans have used, and the southern parts of the Door Peninsula. do use, the quills in artistic embroidery.

318 THE WILD MAMMALS OF WISCONSIN The aesthetic value of the porcupine, dar, white, red and jack pine, and tamarack) which is a most curious creature, is well known and some deciduous trees (willow, beech, in American literature (e.g., Mother various maples, yellow and paper birch, and Westwind’s Stories). It lives in the forests aspens). The porcupine sometimes wanders posing for pictures near resorts, stirring up far after food, into deciduous forests for interest at campsites, and is tolerated in man- leaves (in summer), and often into conifer- aged forests. Local damage is caused in some ous forests where phloem is eaten after mid- tree plantations. October and in winter. In summer, the ani- The status of the porcupine must be mal was observed to graze, so to speak, only watched closely, as it has retreated northward 15-60 minutes per day. Second growth for- significantly in its geography. With land use est is preferred over mature forest (Golley of humankind relentlessly removing trees, we 1957, Brander 1973). can expect less habitat for porcupines. Less In summer porcupines often forage on attention should be given to “control” of a the ground, eating such foods as roots, leaves, “nuisance”, and more attention to preserving and hazelnuts. Porcupines have fed even on an unusual mammal. pond lilies. Bone is often gnawed by porcu- Habitat. The porcupine is found in for- pines, and any trace of salt is eaten. Porcu- ests or open woodlands, particularly the mixed pines have searched human habitations for conifer-hardwoods of Northern Wisconsin and salt (Jackson,1961; Rogers, 1981). Others Upper Michigan (Earle 1978, Kelly 1973, report corn occasionally is damaged. Golley 1957, Dice and Sherman 1922). Reproduction. Shadle (1944,1946, The porcupine prefers for shelter a hol- 1948 and 1951) documented the reproduc- low log or tree, or a small cave. Shapiro tive behavior of the porcupine. Struthers (1949) found dens protecting the porcupines (1928) and others have supplemented this more against wind than cold. Temporary dens information. Breeding takes place from No- may be used, sited as one km away from an- vember to early December. The porcupines other. Fecal droppings signify the presence are promiscuous. The idea of copulation is of a den. Burt (1946) reported an occurrence mind boggling, but presumably a pheromone of 20 porcupines in a winter den. in a urine secretion seduces a female’s co- Foods. Seasonal vegetation includes operation, i.e., holding down her quills. Oth- buds in spring, nuts in autumn, inner bark er observers make no such claims. They men- in winter, and leaves and twigs in summer. tion nose-sniffing and “amorous” noisy vo- The trees utilized for foods are conifers (bal- calizing, sometimes answered with a loud cry sam, hemlock, black and white spruce, ce- of rejection. Betsy Graham, in central Wis- consin, heard the noisy courtship as early as 28 October 2001. Gestation is long (209- 217 days) and the litter size is usually a single young. Occasionally there are two. Birth varies from April to mid-June. At birth the young porcupine weighs about 1.2 lbs, with the minute quills beginning to grow out, all encased in the mother’s amnion. The young is clothed in long, silky hair at birth. It can whine, and has deciduous teeth erupted (Sha- dle and Ploss 1943). Thus, the porcupine is born in a rather precocial state, in a small Quills of porcupine. Showing scales. Po-Chedley. litter, and after a long gestation. In porcu-

TAXONOMIC ACCOUNTS / ORDER RODENTIA 319 pines, which lack post-partum specialization The highest density ever reported for and bonding, intelligence is apparently not porcupines was in northeastern Wisconsin enhanced by this pattern, judging by the lack (Stoeckeler 1950), a high of 96.7 individu- of playfulness and little evidence of intelli- als per square mile of mixed hardwood-hem- gence in young porcupines. lock forest. Golley (1957) suggested upper The young sucks from nipple to nipple. Michigan supported at least two per square Weaning takes place in two weeks. The juve- mile. Brander (1973) reported 40-58 por- niles or young subsist on solid food, and the cupines per square mile in old-growth hard- mother pays little attention to it (Jackson wood-hemlock forest in western Upper 1961). However, Gehring (1993) observed a Michigan, and in Iron County-winter mother defending her young against ravens. counts—at 24 per square mile in cut-over Porcupines are slow to develop sexually. forest areas. In heavily logged areas in sum- Maturity is attained in 1 1/2 - 2 1/2 years. mer, there were 32 per square mile, and Porcupines live about six years in nature, as 43 per square mile in areas selectively long as 10-11 years (Brander, 1971). Aging logged. The mature hardwood-hemlock for- of cementum annuli is reported by Earle and est seemed a winter refuge for porcupines Kramm (1980). Long life seems necessary for on the Upper Peninsula. a mammal with limited fecundity. Remarks. The porcupine defends itself Mortality. There are few enemies of the by turning its rear end toward an attacker. A porcupine. Black bears and fishers occasion- predator is more successful if it attacks the ally kill them, but not often as Seton and oth- face, but the porcupine is quick to turn about. ers have implied. Coyotes eat them. Auto- The tail switches back and forth suddenly driv- mobiles and hunters kill them. A small porcu- ing numerous quills into the attacker’s face. pine may be preyed on by birds of prey, even Although arboreal, the porcupine clam- ravens (Gehring 1993), if the mother does bers best in trees and branches with diame- not defend it. ters less than 6-10 inches. They are too clum- Parasites include a mallophagan, suck- sy to climb trees with great circumference ing lice (Eutrichophilus), ticks (Dermacen- (Harder, 1979). Porcupines are able swim- tor, Ixodes), tapeworms (Shizotaenia), and mers, and although they seldom dig in the roundworms (Dipetalomema, Dirofilaria, ground, they do dig long tunnels in snow ra- Molinema, and Wellcomia) (Jackson 1961; diating away from winter dens. Communica- Doran 1954). Curtis and Kozicky (1944) tion among porcupines is reviewed by Seton counted 1,528 tapeworms and 5,184 round- (1953) and Baker (1983). worms in one female. Porcupines contract Additional Natural History. A mono- mange (Baker 1983). graph on the porcupine was written by U. Home Range and Density. Season and Roze (1989). habitat affect home range (Shapiro 1949, Geographic Variation. There is no geo- Taylor 1935). Porcupines migrate from win- graphical variation in Wisconsin and Michigan. ter to summer ranges, as far as 1500 m Specimens examined. Total, 30. Clark (Dodge 1967). This home range may be called Co.: Wedgee Cr., on Hwy 10, and. Five- a shifting home range. Average daily move- mile Cr. D.O.R. Examined but not pre- ments in Minnesota (Marshall et al. 1962) served. Door Co. Corporate limits Rud- were 393 feet (125 m) in daytime and 264 dville, above canal 1. Near Rock Creek, 1), feet (83 m) at night. In Upper Michigan home Douglas, Forest, Iron, Langlade, Mara- range was a circle about 300 m diameter. This thon, Marinette, Oconto, Oneida, Outag- home range declined; in winter it was restrict- amie, Portage, Price, Rusk, Sawyer, Wash- ed to a circle of 16 m diameter (8 m radius). burn, Waupaca counties.

320 THE WILD MAMMALS OF WISCONSIN ORDER CARNIVORA Family CANIDAE Gray 1821 Meat Eaters Dogs, Wolves, Foxes

The Carnivora is an order of both modern Canids are cursorial, lithe but strong meat- and primitive mammals, descended from an- eaters having large canines, long limbs, long cient Insectivora and their direct descen- ears, and almost a full (primitive) complement dants called Creodonts. Carnivores had spe- of toes and teeth. There are four functional cialized their habits and teeth (early to mid- digits on each forefoot (and a non-functional dle Paleocene epochs) to kill prey. Particu- dew toe as well), four on the hind foot, and larly in the development of the carnassial 42 teeth. The snout and bony rostrum are pair of teeth (P4/ and M/1) and elongation slender, the lower jaw articulates at the men- of the canine teeth is this specialization ev- tal symphysis for nipping and slashing prey. ident (except in some omnivorous Carni- The claws are blunt. Some canids (wolves) vores, in which the carnassials were spe- hunt in packs; but foxes hunt alone or in small cialized for crushing and show little shear). family groups. Seven Recent families are found throughout the world, excepting Australia (which now has introduced the feral dingo and huge Key to the Canidae feral house cats). Some workers include in the Carnivora the pinnipeds (seals, sea li- 1 Skull dog-sized and lightly built, basilar ons, and walruses) evolutionarily derived length (posterior edge of upper incisors from the Carnivora. Five families of carni- along mid-axis to anterior notch of fora- vores occur in Wisconsin. These flesh-eat- men magnum) usually less than 147 mm, ers range in size from the rare, mouse-sized frontal bones concave dorsally “dished”) least weasel (Mustela nivalis) to the huge, ...... 2 omnivorous black bear (Ursus americanus). 1' Skull large and massive, basilar length Most of the predatory, furbearing quadru- usually more than 147 mm, frontals con- peds belong to Carnivora. All are called vex dorsally in profile, never dished .... 3 “carnivores,” a word not to be confused with 2 Tip of tail white, ears black on outer sur- the general meaning of the word, “eaters face, sagittal ridges converge posteriorly of flesh,” which are, of course, described in a V-pattern and they are not conspicu- as “carnivorous”. ous, dentary lacks a prominent step anterior to the angular process ...... Red Fox Vulpes vulpes 2' Tail having a continuous dorsal black longitudinal stripe, ear pinna reddish or rufous, sagittal ridges prominent and bowed laterally, inferior margin of dentary bone having a prominent step ...... Gray Fox Urocyon cinereoargenteus 3 Skull massive with broad rostrum or snout, robust canines (anteroposterior length exceeds 11 mm), upper carnassial exceeds 23 mm, canines extend approximately to and occasionally beyond level of mental foramina, P 4/ - P 4/ Coyote feeding young. By E. R. Kalmbach. breadth (transverse) divided into length

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 321 3,842 $1.40 $5,378 21,800 f pelts may drive numbers up, but low prices not ned,. The value of the harvest is extremely diverse, ranging from t and beaver accounts for aquatic furbearers. $4,278 19,000 $20.88 $18,000 1,586 735 $30.73 $22,500 1,485 3,63127 $12.31 $44,700 $45.47 2,172 $1,000 2,020 61 $11.57 $2,337* 11 2,268 $44.75 $492 83 1998-1999 Total Pelts Value Pelts 1998-1999 Total $1,034,576 million — $14,000 11,469$29,000 524$8,500 $3.23 1,992 $1,692 $2.33 2,778 8,100 $4,500 $1.54 294 ?? $3.26 399 $836 $2.20 9,064 $877 ?? $13.85 $120,000 8,432 5,014 $15.91 $79,000 5280 4,430 $16.49 $73,000 7,374 Harvest1994-1995 Trapped Harvest 1995-1996 Trapped Harvest 1996-1997 Trapped 1997-1998 243,879976 $14.6329335 $3.5 $9.185,6924,660 $43.70 $4.33 $8,9592,334 $1.47 $1,2672,104 $15.63 — $1,450850 $10.38 $8,367 — $72,835 $33.05 — $24,226 — — $2.97 3,304 $69,537 — 618 — 1,773 $2,524 789 27,402 5,337 — 68,230 2,14 9 vest summaries for terrestrial furbearers in Wisconsin. B. Dhuey, personal correspondence. The price o ” Table Car-1. Table Car-1. Table Car-1. Some fur har Table Car-1. Table Car-1.

opossums down, and some furbearers are low, i.e., scarce. Skunks, also opossums, at low prices show many more trapped than skin tremendously high values for raccoons to hardly anything bobcats, weasels, and skunks. The reader is referred the muskra Striped skunk Thousands wasted“Weasels Harvest lowspecies?Trend Opossum Thousands wasted 11,368 2,103 $2.53 $6.74 5.593 $1.53 Bobcat high,trapped high,Pelt harvest value low Fisher DownwardTrend 259 $36.17 4,925 $9,000 51 $27.22 $134,000 862 Species Raccoon Up down UPTrends fox Red down Values Trend 175,973 $8.76 8,662 $1.5 million 119,716 109,417 $8.86 $96,900 89,513 171,255 $14.57 $2.5 million 108,340 Gray fox Down, value alsoTrend Coyote downNumber and Value 1,997 3,800 $9.99 $20.47 $20,000 $78,000 2,072 2,442 752 $9.03 $6,700 1,571 734 $9.37 $6,877 1,120 Gray Fox Coyote Bobcat Fisher Striped Skunk “Weasels” Opossum $20,000 to $23,000. approx. *Coyote value low, Raccoon Fox Red

322 THE WILD MAMMALS OF WISCONSIN from anterior premolar to and includ- and habitats. But they agree in their general form ing last molar is less than 2.7 ...... and characters; the Coyotes everywhere are sons of ...... Timber Wolf* the desert, Ishmaelites living by their wits. Further, Canis lupus they are alike in their vocal gifts—our Ishmaelite is 3' Skull narrow with slender rostrum or also a Troubadour.” — Ernest Thompson Seton snout, canines elongate and less massive (less than 11 mm anteroposterior 1823. Canis latrans Say. In Long. Account of an length), upper carnassial less than 23 expedition to the Rocky Mts... 1:168. Type mm, canines may extend below level of from Engineer Cantonment, 12 mi. SE of the mental foramina of mandible, ratio of P present town of Blair, Nebraska, W Bank of 4/ - P 4/ exceeds 2.7 in maxillary tooth- Missouri River. row ...... Coyote** 1897. Canis pallidus Merriam. Proc. Biol. Soc. Canis latrans Washington, 11:24, type from Johnstown, Nebraska. Not Canis pallidus Cretzchmas, 1826, type from Kardofan, North Africa, now Genus Canis Linnaeus referred to Vulpes. This name pallidus Merri- am is a synonym, and not usually regarded as These are the large dogs in Wisconsin, in- an invalid homonym, which it is and which cluding the domestic dog (Canis familiaris), makes the name unavailable. the timber wolf (C. lupus), and the coyote (C. 1898. Canis nebrascensis Merriam. Science, N.S., latrans). In Wisconsin, some dog X coyote 8:782, a renaming of the homonym Canis hybrids (“coy- dogs”) may be found. I suggest pallidus Merriam. this is seldom.

Canis latrans thamnos Jackson Canis latrans Say Coyote 1949. Canis latrans thamnos Jackson. Proc. Biol. Soc. Washington 62:31, type from Basswood Island, “While we speak broadly of the Coyote as though it Apostle Islands, Ashland Co., Wisconsin. were one species, it should be remembered that scientists recognize at least a dozen kinds that are The name Canis means “dog” in Latin, closely akin, and yet have their own peculiarities and the name latrans means “barking.” The Aztec word coyotyl, from which “coyote” is derived, also means “barking dog.” In early Wisconsin it was called the “prairie wolf” to * Some dogs resemble wolves cranially, but the orbital angle (between tangents to dorsal surface and distinguish it from the gray or northern tim- to the line frontal-top of zygoma) exceeds 52 de- ber wolf, Canis lupus. grees in dogs. Description. Having size of a fairly large ** Coyote X dog hybrids (“coy dogs”), although uncom- domestic dog, the coyote also has about the mon due to low influx of genes already may be chang- same, but slenderer, proportions. The canine ing the characters of the wild coyotes. Gradual introgression eventually breaks down species distinctness. teeth are relatively longer (although there are Many coyote skulls in Wisconsin seem small and have exceptions), and either snout or rostrum of short canines, not extending down to the mental fo- the skull is more slender. The external ears ramina, whereas other skulls are typical of coyotes. are large and pointed. The eyes are usually The coy-dog skull is usually small with short canines, yellowish with a round dog-like pupil. The legs the cranium somewhat arched as in dogs, the rostrum short, and the pelage and body may reflect domestic are long, but the feet small. The forefoot has dog characters. 5 toes, but the innermost is reduced, and each

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 323 hind foot has four toes. The dew-toe on the coyotes (kg ± 1SD) from the years 1978- coyote’s front foot is better developed than 1981, as 12.2-17.2, and for females 9.5- that of the wolf. There is a scent gland at the 13.2. Coyotes older than one year averaged base of the tail. 14.0 ± 1.4 (N = 10) for males, and 11.2 ± The skull is flattened, hardly arched at 0.9 (N = 10) for females. all. The zygomata flare away from the brain- Dental Formula. DF = I 3/3, C 1/1, P case. There is a median sagittal crest, less 4/4, M 2/3 = 42. pronounced than in adult wolves. The pos- Geographic Distribution. Coyotes are torbital processes are large, convex bulges, sparsely distributed but range statewide in instead of being sharply pointed. In a “good” appropriate habitats. The habitats are excep- coyote specimen the canines extend below tionally varied depending on the absence of the mental foramina of the dentaries. This is dogs and wolves to some extent, and are re- one criterion separating coyotes from dogs. lated to year-round food availability. Another character is the high arched crani- Status. Any victimized farmer and his um of the dog. A third is the slender palatal relatives and neighbors will, of course, con- width of the coyotes. The ratio of the length sider occasional depredation as too often. of the upper premolar-molar toothrow to the Appropriate security for the livestock lessens breadth between the alveoli of the first mo- depredation. Coyotes will kill sheep, calves, lars is 3.1 or more. This ratio is 2.7 or even pigs, and chickens, but not as often as gener- less in dogs (Howard 1949). To compare ally believed. They kill fawns, an occasional wolves, see account of Canis lupus. adult deer (usually sickly), and numerous birds, The baculum (62-89 mm in length, Burt, rabbits, and rodents. 1960) is spindle-shaped with a deep trough, Poisoning of coyotes was carried out for i.e., showing a deep urethral groove. Kennel- decades in western states even to the detri- ly (1978) reports the karyotype is 2N = 78, ment of many beneficial fur-bearing mammals. the same as for dogs and wolves. There are To some extent, the known feeding on live- eight mammae. The coyote pelage is grizzled stock carrion led to some wrong assumptions buff to yellowish gray and sometimes rufous and to many false accusations of livestock brown strongly intermixed with black guard predation. A bad reputation exaggerated for hairs. These are dense and coarse, especially the coyote led to intense persecution—hunt- along the mid-dorsum. The fur resembles that ing, trapping, and even bounties in many of the husky dog or timber wolf. The throat and belly is a “dirty” or buffy white. The tip of the tail is blackish. The face, ears, forelegs and feet are brownish, even rust, but around the lips there are whitish hairs. The molt takes place annually, between late spring and autumn, when the fur comes in prime. Pups are yellowish or buff-gray. Male coyotes are larger than females in most measurements. Males often exceed 1,000 mm (39.4 in), ranging up to 1300 mm (51 in) in total length. Skulls average 180 mm (7.1 in) in length. Weights vary to 45 lbs (20.3 kg), and rarely are even heavier in some males. Females seldom exceed 25-28 pounds (11- 12 kg). Smith (1984) lists weights of adult male Skull of Canis latrans.

324 THE WILD MAMMALS OF WISCONSIN states. The coyote seems to be maintaining cluded by timber wolves. The coyote now its genetic identity, increasing its geographic ranges throughout these same forestlands and range, and maintaining its populations in the even crosses ice to visit most of the islands in face of approximately 30-40 percent mortal- Lake Superior and Lake Michigan. It breeds ity caused by humankind. Coyotes feed on on some of them. (It is not a permanent res- harmful mice and rabbits and thereby do a ident on Washington, Rock, or Detroit islands, service. The coyote tends to cull out the starv- but lives on the Door Peninsula.) ing, weak and sickly, benefiting some wild prey The coyote has become more abundant populations. We are skeptical about most in northern than in southern Wisconsin. From claims of coyotes killing adult cattle. St. Croix, Dunn, Eau Claire, Jackson, Mon- Today, trapping is regulated, hunting roe, southern Juneau and Adams, Marque- permitted, and the fur valuable ($10-20 per tte, Winnebago counties and northward along pelt) (Table Car-1). In the not-to-distant past, Lake Michigan up to Brown County, there bounties were paid for Wisconsin coyotes are few coyotes. Rolley reported increases in (Smith, 1984, Table Car-2), a practice that all six state DNR districts for 1991-1992, es- proved both costly and ineffective. pecially in southeast Wisconsin (400 per cent), Historically, the coyote apparently was even 600 percent (Rolley, 1996). The har- unknown in northern Wisconsin and Upper vest for 1999-2000 was estimated as 21,745 Michigan until after 1900 (deVos, 1964). The (Dhuey, 2000), mostly from Grant, Crawford reason for this restriction to the savannas and and Racine counties. A survey of trappers prairies (and avoidance of the northern for- provided an estimated harvest of 10,695. ests) is not clear. Coyotes may have been ex- Coyotes are increasing in the Kettle Moraine State Forest, southeast Wisconsin, and in closely adjacent areas. In the Northwest District re- Table Car-2. Wisconsin coyote harvests or bounty cent sightings are highest in frequency since records (1930-1982) and pelt values. Smith (1984) and 1986. In 2003, coyotes were protected in Dhuey (1999). northern counties during the deer season. 1930 1,949 $5.04 1954 2,557 $1.00 1978 6,788 $47.20 Habitats. Coyotes occur wherever suit- 1931 1,422 3.59 1955 2,118 1.00 1979 2,815 32.18 able prey occurs, but forest edge, clearings, 1932 1,387 4.12 1956 1,785 1.00 1980 2,730 23.70 clear-cuts, farms, and highway rights-of-way 1933 1,785 3.20 1957 no bounties 1981 3,728 26.71 1934 2,111 7.80 1958 no bounties 1982 2,694 19.76 certainly benefit the coyotes. They prefer 1935 2,324 2.14 1959 2,058 1.00 1983 3,238 15.02 mixed woodlands and hardwoods, aspen, and 1936 1,775 4.17 1960 3,673 1.00 1984 2,865 13.44 marshland, but also inhabit unusual (even ur- 1937 1,824 3.36 1961 2,580 — 1985 2,253 14.21 ban) habitats. 1938 1,587 2.37 1962 last bounties 1986 2,535 15.63 A coyote’s burrow may be used for sev- 1939 1,131 — 1963 518 — 1987 2,547 14.53 1940 913 — 1964 — — 1988 797 7.07 eral years. Often the coyote enlarges an aban- 1941 924 — 1965 197 5.21 1989 433 7.06 doned burrow of a badger or woodchuck. 1942 885 — 1966 229 2.22 1990 392 6.17 There may be several entrances for a den. 1943 1,125 — 1967 185 2.38 1991 3,117 5.99 Resting outliers may be used on hillsides in 1944 1,936 — 1968 1,104 5.46 1992 3,554 14.10 clumps of conifers or other trees, even on 1945 4,134 2.32 1969 2,424 5.46 1993 2,727 14.61 1946 3,317 1.09 1970 1,799 5.36 1994 2,442 20.47 snowdrifts. Gier (1968) reported that 10 per- 1947 3,604 2.38 1971 2,715 7.91 1995 3,631 12.51 cent of Kansan natal dens had two females 1948 2,875 1.00 1972 1,849 11.39 1996 2,168 11.00 and double litters. Camenzind (1978) and 1949 3,135 1.00 1973 3,839 17.25 1997 3,330 10.00 Nellis and Keith (1976) found den sharing to 1950 2,844 1.00 1974 2,753 17.25 1998 2,092 7.06 be rare. W. E. Berg (personal correspondence) 1951 2,826 1.00 1975 3,153 18.81 1999 ?? ?? 1952 2,638 1.00 1976 4,986 33.99 2000 ?? ?? doubts if such sharing occurs in Lake States 1953 1,696 1.00 1977 5,125 29.44 2001 ?? ?? coyotes. The nest is often bare earth in a bur-

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 325 row. Long (1964) reported one made above ground and the high water table in a Kansas marsh. Tunnels usually run 3-4 feet below the ground surface. Foods. Food studies include analysis of stomach contents (Sperry 1941, and others), scat analysis (Beckoff 1978), and following trails in the snow (Beckoff 1978, Berg and Chesness 1978). Mice were the primary food, and deer carrion and birds were high in incidence. Excluding livestock carrion, the snowshoe hare, shrews, and the porcupine were often eaten. In a national study, Sperry found

Maps showing geographic distribution of Canis latrans in Wisconsin and North America.

326 THE WILD MAMMALS OF WISCONSIN rabbits and hares most important as prey, such things as berries to carrion, reflecting mice important as well, while deer remains availability and season. In southern Wiscon- and other carrion were often scavenged. sin livestock carrion is 44 percent and rabbits Comparing the main foods of coyotes in 21 percent. Some vegetation, birds, and oth- Minnesota and Wisconsin, the patterns are er foods are eaten (Fruth, 1986; Niebauer and similar. Smith, in 1978-1979, examined Rongstad, 1977). 8,674 scats of coyotes in northwest Wiscon- In Minnesota, the 1,558 aforementioned sin, also examining over 1500 stomachs from stomachs collected over an 8-year period northern Minnesota, not so many in Wiscon- showed that white-tailed deer meat was the sin. White-tailed deer flesh is fed on through- predominant food, occurring there in 48.6 out the year, but not much from August-No- percent of stomachs (41.8 percent total vember in Wisconsin, or May to August in weight of foods). In that study, livestock eat- Minnesota. In Wisconsin, deer as food peaks en as carrion occurred in 18.9 percent (23.3 in December and continues through June. percent total weight). Mice were found in 23.1 Snowshoe hares are fed on year round, but percent (only 4 percent total weight). Porcu- the numbers diminish in winter. The number pine was found in 20.1 percent (4.0 percent), of hares eaten is correlated with the hare cy- and snowshoe hare in 18.4 percent (12.8 cle. When hares are scarce, more porcupine percent). These food items were 85.5 per- flesh is eaten (Smith, 1984). cent of the total contents. A study of 670 In Wisconsin the percent frequency of coyote scats collected in Minnesota over six small mammals eaten (up to 19 percent), of- years showed deer in 40.9 percent, with low ten Peromyscus leucopus and Microtus penn- frequencies in August and October. The deer sylvanicus, rises in September, October, and remains were over 54.8 percent in winter and November, when mouse populations are high. only 27.1 percent in snow-free woods. A dif- Although the frequency of rodents eaten is ference found in study of scats compared to high, the weight is low, about 4 percent by other analyses of coyote diets was a signifi- weight. Some vegetation is eaten year round, cant amount of plant parts and seeds. Data but chiefly from June to October. Vegetal foods from some coyote trails were summarized by include berries, buds, grasses and seeds. Berg and Chesness (1978). Baker (1983) lists There was noted correlation between fat numerous plants and other foods of the coy- indices in coyotes and deer flesh in the diet. ote, in Michigan. Coyotes that fed on deer fed seldom on live- Some deer predation may result from stock. In Minnesota, and not in Wisconsin, pack hunting (Fruth, 1986). About 14 per- the percent of lifestock was 5 percent over cent of capture attempts on deer were suc- 11 years. Ruffed grouse were taken during cessful. A deer was caught by a pack of coy- their spring drumming. Fawns as prey peaked otes after a chase of 2,254 m (1.4 mi.), and in May-June, and declined in July (Smith, another after a chase of only 15 m (15 yds). 1984) and thereafter. The diet varies from Both were fawns. On 703 km of coyote trails coyotes fed on deer 54 times; 52 times carrion was involved (Smith, 1984). Reproduction. Coyotes are monestrous, resembling the domestic dog in reproductive pattern (Hamlett 1938, Gier 1968). Produc- ing a single litter per year generally leads to monogamous parents. They produce large litters and care for the pups. Coyotes may pair Coyote. Scratchboard, by Dan Metz. for several years, but not necessarily for life.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 327 Juveniles may breed their first year if envi- fur, kill them as “livestock killers,” and occa- ronmental conditions are favorable (Gier sionally run over them with automobiles. Poi- 1968). Usually coyotes commence breeding son baits were used extensively in the past, in their second year. In northern Wisconsin but are illegal today. counties juvenal breeding is unlikely. Smith Gier et al. (1978), Pence and Custer (1984) reported pregnancy rates in juveniles (1981), Custer and Pence (1981), and Voigt up to 23-35 percent. Adult females showed and Berg (1987) reported on parasites and 58 percent pregnant, and yearlings 38-40 diseases in coyotes. The tabulation of Gier et percent. Not all the adult females breed. al. is more than a species list (Table Car-3) Breeding occurs between February and for the effects of a parasite or disease “in March. Estrus lasts 4-5 days. Texas cannot be interpolated to Nebraska. . Ovulation is late during the estrus period .Neither can effects... during the winter be (the day varies). Food availability directly in- transferred to summer... or those during times creases corpora lutea counts (4.5 to 6.5 cor- of nutritional stress be transferred to times of pora lutea), and 50 to 80 percent of females plenty.” Sometimes lice and mange mites are are ovulating females. Smith (1984) found 5.5 tolerated by coyotes, and then suddenly the ova to produce 4.4-4.8 embryos in Wiscon- complex of the two is a killer. Lists of para- sin. Transuterine migration of embryos is com- sites are given by Jackson (1961) and Baker mon, leading to approximately equal implan- (1983). Some diseases include parvovirus, tations in each uterine horn. Gestation lasts canine hepatitis, rabies (rarely), plague, and 60-63 days (Kennelly, 1978). especially distemper. Up to seven pups are born, in early Home Range and Density. Following 217 spring. The young weigh about 250 g at birth. winter trails in Wisconsin, Smith (1984) found Their eyes are closed, but the skin is furred. 61 percent of coyotes solitary, 32 percent in The eyes open and the ears become erect in pairs, and only 6 percent in groups of three. 10 days. The young emerge to play after 3-4 Huegel (1979) found as many as five coyotes weeks. Both parents may bring food to the in a pack; whereas pairs comprised 87 per- young, although in Wisconsin usually the cent. Although family groups are typical of mother alone bring dead rodents or regurgi- coyotes, they seldom form packs as do other tated foods. Weaning occurs in about 8 weeks. wild dogs. Fruth (1986) gives the home range The mother teaches the pups to hunt. In nine for Wisconsin coyotes as 5-10 mi2. Smith months the pups attain adult size. (1984) calculated home range for males as Aging of coyotes has been studied by 22.6 km2 and for females 15.6 km2. Lindhart and Knowlton (1967) and Roberts Smith (1984) found that Juveniles com- (1978). Juvenile survival is lowest in mid-win- prised 80 percent of the dispersers. Year- ter (Smith, 1984). The mean adult survival lings comprised nine percent. Nomadic wan- rate was 62 percent (SD = 16). Coyotes may derings accounted for 11 percent. One ju- live nearly 16 years (Crandall, 1964). venile dispersed 160 km (100 mi.), and an Mortality. Golden eagles, rare in Wiscon- adult female moved 440 km (275 mi.) into sin, have attacked adult coyotes. Badgers have central Iowa. eaten the pups. Wolves might drive away or Home range determined for 100 Minne- kill an occasional coyote. As wolves return to sota coyotes from 1970-1975 varied by age Minnesota and Wisconsin, this interspecific and sex. Adult males averaged 68 km2, and aggression will doubtless increase. Starvation for adult females the average was 16 km2. in coyotes may cause some mortality. Human- Males overlapped and wandered freely, but kind and disease are the worse enemies of females excluded one another from territories. coyotes. Humans hunt them, trap coyotes for Of their total home ranges females occupied

328 THE WILD MAMMALS OF WISCONSIN Table Car-3. Some parasite diseases affecting Wisconsin coyotes. From the table in Gier et al., 1978, Pence and Custer, 1981, Trainer and Hale.

Parasite Geographic Range Normal Host or Source Demodectic Mange The mite Demodex canis Cosmpolitan in dogs, wolves, foxes Sarcoptic Mange The mite Sarcoptes scabei Dogs, wolves, foxes Wisconsin Ixodes kingi Raccoons, foxes Dermacentor andersoni Mammals Trichodectes canis Canids North America Pulex simulans Carnivores North America Rabies Rhabdovirus Cosmopolitan in canids Distemper Virus of Carve Saliva, feces Cosmopolitan Hepatitus (fox encephalitis) Adenovirus Saliva, feces Cosmopolitan Histoplasmosis Histoplasma Inhalation Cosmopolitan Musty river bottoms Toxoplasmosis Toxoplasma Feces, urine Cosmolitan dogs, cats

54, 22, and 61 percent in pre-denning, den- Coyotes crossmate with dogs to produce ning, and post-denning periods. More than 70 fertile coy-dogs, which show intermediacy of percent of young-of-the-year disperse from characters (Seton, 1929, Kennelly and Rob- their natal territories. Dispersal may be as far erts, 1969, Mengel, 1971, Gipson, 1978). as 100 miles (160 km), but usually 2-4 km Coyotes also rarely cross with wolves (Kole- (Beckoff 1982). Prior to dispersal juveniles nosky, 1971). An emerging problem is hy- average square 5-8 km2 in their wanderings. bridization with domestic dogs, which are in- In Upper Michigan, density of coyotes creasingly in contact with coyotes. The coy- in Alger and Schoolcraft counties was deter- ote conceivably could be genetically altered mined as one per 4 miles (6.4 km) by follow- out of existence. It is somewhat difficult to ing snow trails (Ozoga and Harger 1966). find a good skull of a coyote in some places Seasonal changes were reported by Andert in Wisconsin, a skull showing the coyote’s and Gipson (1979). pure characters. Nationally, the coyote seems The relation of age structure in a popu- to be maintaining its form, behavior, and in- lation for coyote, wolf and both foxes is dis- creasing geographic range. Gipson et al. cussed by Kleiman and Brady (1978). Smith (1975) found decreased fecundity in the hy- (1984) obtained similar results in winter pop- brids. The heterozygosity of Wisconsin populations, with 50 percent in juveniles, 20-37 ulations should be appraised. In the UW-Mad- percent yearlings, and a few examples of old- ison collection are 3 coy-dogs from Fort Mc- er age up to 12.5 years (Car-4). Coy and one from Douglas County (3 mi. N, Remarks. Coyote communication, 9 mi. W Solon Springs). which is complex and varied, is reviewed by Lehner (1978). The yipping, barking, and howling constitute some of nature’s most Table Car-4. Mortality Table in Minnesota (1970-1982) lovely nocturnal music. Coyotes also com- and Wisconsin (1978-1983). After Smith, 1984. municate by scents and fecal sign. The intelligence and adaptability of coyotes are leg- Minnesota Wisconsin endary. A review of coyote-damage and con- Trapped 37 22 trol research is given by Sterner and Shu- Shot 27 50 make (1978) and by Wade (1978). Econom- Snared 11 — ic importance of the coyote, a long standing Car killed 6 12 Dog killed 0 0 hot issue, is ably discussed by Baker (1983) Natural causes 6 6 and Voigt and Berg (1987). Other 12 10

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 329 Smith (1984) reported “Sampson” (lack- northern Florida, all assigned to one race char- ing guard hairs) coyotes (in both sexes) from acterized by grizzled gray color and rather 1979-1981. They were produced by a pair small size. This wide-ranging race was known of normal appearing coyotes. as Canis lupus lycaon Schreber. Recently Additional Natural History. Voigt and Nowak (1996) ascribed Wisconsin and its sur- Berg (1987) mentioned that even with thou- rounding regions to a wide-ranging race Ca- sands of references, the biology of the coy- nis lupus nubilis Say, with type from Iowa. ote is an enigma today. Bekoff (1977) is a The wolf was formerly distributed fine reference source. Three fine works are throughout much of Eurasia and North Amer- Young and Jackson, 1951; Beckoff, 1978; ica, and in this hemisphere ranged from the and Dolnick et al., 1976. Arctic slope and the coasts of Greenland, Geographic Variation. There is a single occurring on many islands, and extending subspecies found in Wisconsin and Michigan. deeply into southern Mexico (see Map). Now Specimens examined. Total, 16. Adams, they are greatly restricted by eradication in Bayfield, Columbia, Florence, Iron, Mari- many places, by prolonged poisoning of pred- nette, Oneida, Portage, Price, Rusk, Saw- ators even until the1950’s, and by over-hunt- yer, Other records in Wisconsin (Balliett and ing and trapping. The intolerance of people, Taft, 1978): Winnebago Co. Michigan. Del- as well as removal of habitats and routes of ta Co.: Big Summer Island 1. dispersal, led to the restriction of wolves to a few states along the Canadian border, much of Canada, and Alaska (in North America). Canis lupus Linnaeus Presently the Timber Wolf is found chiefly Timber Wolf or Gray Wolf along the northern border of the contiguous states, where it is classified as endangered or “Those Wisconsinites who cherish this state’s threatened. It is abundant in northern Cana- history, its abundance of natural resources, and its da and Alaska, and reintroduced to Arizona. leading role in environmental awareness must not When wolves are downlisted from endan- allow the timber wolf to vanish a second time, gered to threatened, problem wolves that because this state may never get another chance.” destroy pets or livestock may be removed. — Dick Thiel, The Timber Wolf in Wisconsin After wolves fill up remote habitats, any trans- location of wolves to such places becomes 1758. Canis lupus Linnaeus. Systema naturae, 10th impractical, and even results in their extermi- ed., p. 39. Type from Sweden. nation. The only practical removal seems selective and highly restrictive hunting or trap- The word Canis means dog, and lu- ping (?by special conservation agents). pus means wolf. Eastern wolves in North America are more likely to be called Tim- ber Wolf rather than Gray Wolf, but both names are popular. Taxonomically revised by Goldman in Young and Goldman, 1944. This old-fash- ioned appraisal, describing microgeographic variants as races, surprisingly refers wolves from a vast eastern region ranging from Min- nesota, upper Michigan, and Wisconsin, northwards to Hudson Bay, into Ontario and Quebec, and southeastward to Ohio, even Timber Wolf. By E. Hamerstrom Paulson.

330 THE WILD MAMMALS OF WISCONSIN Canis lupus nubilis Say coyotes (Chiarelli 1975). The canines are not Timber Wolf or Gray Wolf widely separated as in dogs. Across the upper molars the breadth divided by the width 1775. Canis lycaon Schreber. Die Saugthiere... Theil of the last molar is 5.4 or less, whereas it is 2, Left 13, pl. 89. Type from Quebec, Quebec 5.5 or more in dogs. The occiput of the wolf (type locality fixed by Goldman, J. Mamm., is developed for the origin of powerful jaw 18:38, 1937), muscles, and in the adults a large sagittal-crest 1937. Canis lupus lycaon: Goldman. J. Mamm., develops. It extends posteriorly above and 18:45. parallel to the flattened braincase, not dip- 1823. Canis nubilis Say. Long’s expedition... Rocky ping ventrally as in other canids. No other Mountains. I. P. 169, Type from Council Bluffs, Wisconsin canid has a canine 13 mm or more Iowa. See also Cory, 1912: 321. across the base. 1996. Canis lupus nubilis: Nowak. Canadian Cir- The baculum is trough-shaped and large. cumpolar Inst. pp.387, 395 map. For 1995. It bows upward, and the broad groove extends to 4/5 of the length. The length varies from Description. The largest wild dog in Wis- approximately 83-99 mm (Burt, 1960). There consin, the massive size and coloration of the are 10 mammae, 5 on each side. Small anal wolf are similar to that of the Husky breed of glands are present. There are 2N = 78 chro- domestic dog. The skull is massive, about 230- mosomes, as in the coyote and domestic dog 260 mm (10-11 in.) long and 120-140 mm (Chiarelli 1975). (6-7.25 in.) in zygomatic breadth. The fur is coarse, almost shaggy, with The appearance of a wolf suggests to long, black-tipped guard hairs and short, dense many people the word “powerful”. The ears grayish or brown underfur, which shows are long but less pointed than in the coyote, through as the fur ripples with movement. The the body is long with elongate, slender legs guard hair has a black tip, a cinnamon or buff having large feet, the heel pad of the front subterminal segment, and it is gray basally. foot at least 4 cm (1.6 inches) wide, armed Various pelages show yellowish, rust, iron with robust, blunt claws on the four toes (also, gray, or mostly black. The tail is blacker at its there is one small dew-toe). There are four toes tip. The belly and inner sides of the legs are on each hind foot. There is a bristly-haired scent whitish, grayish, or buff, and the underside of gland at the base of the tail, never found in the tail tends to buff or gray (less black). Mid- domestic dogs, a large, hairy “ruff” on the dorsally the concentration of black usually cheeks, a wide nosepad, and eyes with a round intensifies. Cinnamon or reddish brown tones pupil. The wolf often runs with its brush-like tail “streaming behind,” whereas the tail of the coyote is more often held low. This behavior varies depending on the social status of the animal (both in wolf and coyote). The skull is huge with robust teeth. The cranium, not including the sagittal crest, is flattened so that the orbital angle (from frontal down to zygoma) is 40o to 45o compared to more than 50o in wolf-like dogs (Mech, 1974; Iljin, 1941). The canines are not elongate, but may extend down to the level of the mental formanina in some wolves. The muz- Orbital angle of wolf skull, which is only 40-45°, less zle and rostrum are relatively wider than in than in dogs. Ilgin (1941).

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 331 show on the grayish legs, and are intermixed size (Wydeven et al., 1993, 1995, 1999 etc.). on the crown (especially on the cheeks), and Mid-winter populations ranged from as low about the ears. Wolves vary in the intermix- as 15 wolves in 1985, to as high as 200 or ture of colors. Some are melanistic, white, or more wolves in 1999 (Wydeven et al., 1995, reddish-brown, but black and white phases are 1999, 2002). The minimal wolf population very rare in the Great Lakes region (Kolenos- in 2005-6, was 415-502 wolves, in 115 ky and Standfield, 1975). The foot pads are packs, known from at least 29 counties. black. The fur molts annually in late spring and I was interested in depredations, espe- summer, over a period of several weeks (Young cially angering many hunters recently by kill- and Goldman 1944). Prime fur is attained by ing their dogs. Depredations have been grad- growth of summer fur into winter (Stains 1979). ually increasing in incidence year after year, Tufts of hair help protect the footpads in win- and in 2002 consisted of calves, two cows, a ter. The young molt juvenile fur and replace it horse and its colt, some sheep, deer on game with adult fur in about 30 days. farms, and attacks on about ten domestic The average total length is about 160 dogs. Obviously, considering both the pros cm (63 inches), up to 175 cm (about 69 in.), and cons, the progress reports of the Wis- and weights vary from 23-41 kg (50-90 lbs). consin Department of Natural Resources on A few weigh as much as 45 kg (100 lbs.). our wolves seem fair and honest, and that is The mean weight of adults live captured in essential for management of a rather contro- Wisconsin was 28 kg (62 lbs) for 13 females versial species, which seems now to be ac- and 35 kg (77 lbs) for 18 males (Wydeven et cepted as a valued carnivore in Wisconsin. al. 1995). Males are larger than females. Win- ter wolves weigh more than summer wolves. See Description above. Dental Formula. DF = I 3/3, C 1/1, P 4/4, M 2/3 = 42. Distribution. The wolf formerly ranged throughout Wisconsin and surrounding states. Now it is restricted to a few counties (a recent report lists 11) in northern Wisconsin. By 1995, it became re-established in the Central Forest (this forestland, a useful and somewhat distinctive deer management region, is discussed in the account of the white-tailed deer beyond). Some wolves move back and forth across the Minnesota boundary, in northeast Minnesota and northwest Wisconsin. The present-day population straddles the Michi- gan-Wisconsin boundary. Status. Since 1974, the timber wolf in the Great Lakes region and northeastern United States has been listed as an endangered species by the federal government. The wolf soon (2007) may be downlisted to threatened status. The Wisconsin DNR classified wolves as endangered in 1975, and since Skull of Canis lupus. Bee and Hall (1956), courtesy of 1979 has annually estimated the population James Bee.

332 THE WILD MAMMALS OF WISCONSIN In Upper Michigan, the gray wolf thrived until white humans eradicated most of them in the late 1800’s. A population of about 100 wolves persisted until 1955 (Arnold 1955). Bounties caused the numbers to dwin- dle thereafter, to seven in 1958, and only one wolf was bountied in 1959. The bounty law was repealed in 1960, and by 1965 full protection was accorded the wolf. In the 1960’s the population was reported as perhaps 12-25 wolves, with sightings reported in nine Michigan counties. By the late 1960’s the Upper Michigan wolves were nearly extirpated. After the population fell to six (Hen- drickson et al. 1975) four radio-collared wolves were transplanted from north-central Minnesota (Weise et al., 1975). They died within eight months. North American recent distribution of timber wolf.

Map showing recent geographic distribution of Canis lupus in Wisconsin. Formerly widespread in North America, but resticted in recent years to the Northern Rockies, northern Wisconsin and Minnesota, New England, and perhaps Arizona. After Wydeven and others. ~ 1995-2006.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 333 In 1961, Jackson had reported that the ward. Perhaps 80 existed then in Upper Mich- wolf was “making its last stand in the United igan. Wolves now have returned to some for- States in the northern parts of Michigan, Wis- ested counties in central Wisconsin. Educa- consin, and Minnesota”, but it had little chance tion programs have apparently taught peo- of survival in Wisconsin where its numbers ple about the value of wolf preservation. were reduced “probably to less than 50.” He Heavy fines for poachers also may have mapped records of wolves in almost all the helped the wolves (Table Car-5,6). Wisconsin counties prior to 1900 (as early as Wisconsin wolves varied from a low of 1827 in Manitowoc County), but in few coun- 15 in 1985, to 40 in 1991. The mean annu- ties after 1900. In 1914 or 1916 (see Thiel, al increase was 1.21 from 1986-1991. In the 1993), one was killed in Wautoma, in central early years of recovery human-caused mor- Wisconsin. Other reports, since the turn of tality was intense, but it has lessened. Canine the century, range from Burnett and Barron parvovirus was prevalent in the mid 1980’s counties westward, Oneida (1930’s), and probably caused high mortality. This prob- Chippewa, Marathon, Shawano, Brown, and lem too has diminished (Wydeven et al., 1995; Door counties eastward. Wolves were present Wydeven, 1993). Recolonization from Min- occasionally on Washington Island and in the nesota continues (Wydeven 1993), and the Apostles. Even in northern Wisconsin wolves habitats available suggest the wolf population dropped from sight, reoccupied counties in will continue to increase both in Wisconsin northern and central Wisconsin (see Map), and and Upper Michigan, on public and private lone wolves have occurred as far south as lands. The landscape (or environment) seems Portage in Columbia County (Mech et al., favorable for more Wisconsin wolves, judg- 1995). The population is recovering in Wis- ing from an analysis by Mlandenolf et al. consin, and also has re-established itself in (1995), relying on fractal analysis of patchy Upper Michigan. vegetation, road systems, and other factors. Jackson (1961) had estimated the wolf The steady observed increase and spread of numbers in 1835 as 20,000-25,000. [Wyde- wolves is likewise indicative of future success. ven (1993) suggested 3,000-5,000 wolves was Regulated hunting perhaps will be allowed in a more accurate estimate.] Human persecu- Wisconsin when the wolves exceed 300 (see tion, loss of habitat, and reduction of ungu- table Car-6). lates had already eliminated wolves from south- The Wisconsin Department of Natural ern and central Wisconsin by 1900. The bounty Resources has recently published in the news system, from 1865-1957, was a major factor media a concise history of wolves, succinct in extermination of the wolf. By the 1940’s, and interesting, as follows: 1830’s, a state- fewer then 100 wolves remained, and by 1950 wide population of 3,000–5,000 wolves; there were fewer then 50 (Thiel, 1993). In bounties offered in 1865; no wolves remain- 1957, the wolf was protected, but probably ing in the southern two-thirds of Wisconsin became extinct in that decade (Thiel, 1978). by 1900; wolf bounties were lifted, after mil- By the 1970’s wolves were recolonizing Wis- lions of dollars were spent to eradicate this consin from Minnesota (Thiel and Welch, carnivore, 1957; federal protection as an 1981). Thiel (1993) recounted the reappear- endangered species, 1974; the Wisconsin ance of wolves in Douglas County (in 1975). DNR declares our wolves endanged [and few The slow re-population in northern Wis- if any remained in Wisconsin] 1975; five wolf consin has continued to this day. Most killed packs were found in Wisconsin numbering are shot or killed by cars. At least 83 (in 1995) about 28 individuals, 1979; the federal gov- wolves were found in northern and central ernment reclassifies gray wolves in Wiscon- Wisconsin, and wanderers may range south- sin from endangered to threatened, and there

334 THE WILD MAMMALS OF WISCONSIN were about 323 wolves in Wisconsin, March rock crevices, and other protected sites. A natal 18, 2003. Note that the wolf is still protected den usually has a single entrance, but occa- and managed in Wisconsin under “threat- sionally there are two. The diameter of the ened” classification. entrance is about 36-63 cm (14-25 inches). Although wolf furs are popular for trim No mention is made of an internal nest. A den on coats and as wall hangings, there is no may be re-used for several years. Den-site se- commerce allowed at present in Wisconsin lection is discussed by Unger (1999). In winter because of the wolf’s potected status. Fur wolves may burrow into snowbanks, under fall- harvests are permitted in Canada and Alaska en trees, or even sleep in old beaver lodges. (Carbyn, 1987). They may endure a blizzard by lying in the snow In literature, the wolf has an awesome curling the tail over forepaws and nose. reputation of following sleighs, attacking chil- Foods. Wolves are opportunistic preda- dren, and so on. These accounts are largely tors that generally specialize on ungulates. The fictitious, and there is only one record of North white-tailed deer is the chief food for wolves American wolves attacking a man (Peterson, in Wisconsin (Thiel 1993; Thompson 1952) 1947). (In 1994, news accounts reported that and Upper Michigan (Baker 1983). The wolf a Wisconsin child lost an arm to two caged pack succeeds in running down and killing wolves in a northern Wisconsin zoo.) Never- young, old, weak, or sickly individuals (Fuller theless, when wolves are more numerous and and Keith 1980). After 1,000 m, the chase is come into increasing contact with people, usually abandoned. A large carcass, such as a common sense and responsible parenting are moose, allows the pack to gorge and re-group necessary. Always ensure the security of chil- (Mech 1966). Some wolves eat up to an esti- dren in remote areas. [Virtually any wild mam- mated 9 kg (20 lbs.) of flesh each. Wolves mal can be dangerous to man in certain cir- normally can go 3-4 days without food, but cumstances.] Livestock also requires security, have been known to go 7 -17 days without and there will be some depredations of un- eating (Paradiso and Nowak, 1982). A healthy protected domestic animals. wolf needs a mininum of 1.7 kg (3.5 pounds) Habitat. Forests are the home for Wis- of flesh per day. Many studies on feeding of consin wolves. Presently the mixed conifer- timber wolves have been made by Mech and hardwood forests and forested wetlands of his associates (1970, 1974, 1977, 1982; northern Wisconsin are the most prevalent Mech and Frenzel 1971; and others). Other habitats used by the wolves (Mladenoff et al. foods used by wolves in in Upper Michigan 1995). Wolves do not avoid open country and include shrews, snowshoe hare, red squirrel, will wander through many habitats. Tradition- deer mice, meadow voles, jumping mice, an ally wolves only occurred in the most remote occasional porcupine, ruffed grouse, crayfish, regions were they where free of human per- and grasses (Manville 1948, Stebler 1944, secution. In recent years wolves have moved 1951). To this list may be added raccoon, into farmlands. The major factors determin- woodchuck, flying squirrel, chipmunk, red- ing suitable habitat are lack of human perse- backed vole, bog lemming, muskrat (Baker cution and presence of adequate prey (e.g., 1983), various plants, feral cats, some livestock, white-tailed deer and beaver). and carrion. A wolf in Wisconsin was even A den sketch is provided in Carbyn (1987). known to feed on fish and great blue heron The natal den is dug at a well-drained site usu- (Ardea herodias) chicks at a heron rookery ally on a ridge, by the parents, or is enlarged (Robinson et al., 1991). Mandernack (1983) from an abandoned woodchuck, fox, or bad- discussed food habits of Wisconsin wolves. ger hole (Young and Goldman 1944). Occa- When does are solitary, or when does sionally they use a beaver lodge, hollow logs, are with fawns, wolves often hunt them alone

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 335 or in pairs over wide regions. In winter when months, after which they are moved to some the deer yard together, wolf packs move from outlier (often called a rendezvous site), remain- yard to yard preying especially on old deer ing until late summer or early fall (Fuller, and on the numerous young. Many other ani- 1989). The same den may be re-used. mals scavenge on the remains of wolf kills in- The pups are born furred, eyes closed, cluding foxes, bears, coyotes, and mustelids. ears non-erect and closed, with the fur col- Small rodents gnaw the bones. Birds scaveng- ored a “sooty” drab whitish color (Jackson ing at wolf kills include eagles, ravens, crows, 1961, Mech 1970). Weighing about a pound jays, chickadees, and even woodpeckers. (0.7 kg) each, the growth is rapid (Van Bal- Reproduction. In a typical pack hierar- lenberghe and Mech 1975). The eyes open chy only the socially dominant (an alpha) fe- in 11-15 days (Mech 1970) and the ears stand male produces pups. They are sired by the erect about the same time. The pups begin to dominant male. The other members of the spend time outside the natal den at 10-24 pack assist in caring for the pups and the at- days, but normally remain near the den site tentive mother, thereby protecting their own for another month (Fuller, 1989) unless the genes (by kinship), and also performing a de- mother moves them. Pups are moved about fense of the pack territory. When prey is abun- by the mother when 8 to 10 weeks old, usu- dant, two females may produce litters in the ally in June (Mech, 1970). They are weaned same pack. If a subordinate mates, it may be by 50 days, and pack members regurgitate chased out of the pack (Peterson 1979), but meat for the pups to eat during most of the may bring forth pups. summer. Another female may occasionally The female is monestrous, and rarely is “babysit” the pups. The mother may move capable of conception at 10 months (Medjo the young to as many as 10 outliers at this and Mech 1976). The reproductive pattern is time of development. By late summer the pups similar to that in dogs. Usually breeding does (2/3 grown) join the pack in their nomadic trav- not commence until 22 months, and often does els. Milk teeth are replaced by 16-26 weeks not occur until the third year. Wolves may breed (Schonberner 1965). Aging of wolves by ce- at least until their 11th year if they live that mentum layers has not been successful. Wolves long. They probably may live occasionally to live at least 15-16 years (Crandall, 1964). 16 years of age. Under intense exploitation of Mortality. The chief mortality is caused wolves, as many as 90 percent of adult females by humans by illegal shooting and trapping will breed, but in unexploited populations only and by automobiles. Human intolerance and about 60-65 percent breed. human activities helped to eradicate wolves, The female has estrus about five to sev- but changing social attitudes have been in- en days in late winter. Mating begins in Feb- strumental in protecting the species. Fortu- ruary or earlier. Copualtion may last over 30 nately for the Wisconsin wolves, human- minutes, and the pair may become “locked” caused mortality declined from 72% from as in domestic dogs. Wolves court much as 1979-1985 to 22% in 1986-1992 (Wydeven, do dogs, but remain paired for life (Mech 1993). The Wisconsin Department of Natu- 1966, 1974). Gestation is about 63 days. The ral Resources is studying the impact of the litter size is about 5-7 pups (varying from 4- northern highway 53 on Wisconsin wolves 10 or more, or about 6, 1-11 according to (Shelly and Anderson, 1995, unpublished). Paradiso and Nowak 1982). The pups are Under natural conditions starvation and born in mid-April in a specially excavated natal exertion lead to higher mortality rates when den or at some alternate site prepared sever- prey species such as deer or bison become al weeks in advance of the births. The pups low in density. Mech (1977) found that low are kept at the den area for the first two food availability caused starvation in pups,

336 THE WILD MAMMALS OF WISCONSIN reduced litter size, and increased intra-pack Mech 1991). Densities varied from 1/23-111 aggression. Mortality increased among the km2.. Thiel (1993) reports home range and wolves, and greater numbers became “lone” densities in Wisconsin, from 90-300 square wolves. A cougar or bear might prey on miles, with densities varying from from 1/ 29- wolves, but the wolves may occasionally kill a 93 mi2. In 1990-1991, there were 12 packs, cougar or bear. with pack size averaging 3.2 wolves, territory Wolves harbor rabies virus, tularemia, 2,000 km2, and a wolf density of 18.3/1,000 tapeworms, spiny-headed worms, round- km2 (Table Car-5,7). worms, fleas, ticks, and lice as do other canids Home range on Isle Royale of a winter (Mech 1970; Mech, Thiel, Fritts and Berg, wolf pack was about 272 square km (105 1985). Especially serious are sarcoptic mange, square mi). In northeastern Minnesota, Mech blastomycosis (Thiel et al., 1987), distemper, et al. (1971) and Van Ballenberghe et al. and probably parvovirus (in the 1980’s, see (1975) found the home range to vary from Wydeven, 1993). Wolves carry ticks that har- 20-56 square miles. During mid-summer the bor Lyme disease (Thieking et al., 1992). In home range of an adult female with pups was Wisconsin some internal parasites include 18 square km (7 square mi ) in Ontario (Kole- tapeworms, hookworms, Giardia, Entamoe- nosky and Johnston, 1967). ba, Alaria, and Capillaria (Archer et al., The territory is defended by scent mark- 1986). Starvation and intraspecific strife cause ing (Peters and Mech 1975, Rothman and some mortality. Mech 1978), howling (Theberge and Falls Home Range and Density. Some densi- 1967), or active aggression toward intruders ties and home range data provided by Car- (Mech 1977a). The howls serve to call the byn (1987) and Wydeven et al. (1995) indi- members together and to warn intruders to stay cate the timber wolf home range areas vary a lot, from a mean annual home range of 146 km2 – 349 km2 in northern Minnesota, to 253 Table Car-6. Recovery of Canis lupus in Wisconsin. After km2 in central Manitoba, and 179 km2 in Thiel, 1993; Wydeven et al. (1995); and Wydeven (1995). There were 18 packs. After 2000, there seem over double this northern Wisconsin. From very small summer number. home ranges, Carbyn reported a radio-collared female that from October to March Year Locality Number 2 wandered over a remarkably large 2,819 km 1975 August 3 Douglas Co. 1 area (1,088 miles2), entering six pack territo- By 1979 Douglas Co. 4 ries. Young adult wolves, usually as yearlings, 1979 July 9 Lincoln Co. 1 may disperse from their natal pack to find a November Lincoln Co. 1 1980 N Wisconsin 30 mate and establish a new territory, or join an 1980’s N Wisconsin approx 50 existing pack. Some wolves may remain with 1981-1982 N Wisconsin 27 the pack throughout their lifetime (Gese and 1982-1983 N Wisconsin 19 1983-1984 N Wisconsin 17 1984-1985 N Wisconsin 15 Table Car-5. Timber wolf mean annual home range and 1985-1986 N Wisconsin 16 winter density in central North America (modified from Car- 1986-1987 N Wisconsin 18 byn 1987 and Wydeven et al. 1995). 1987-1988 N Wisconsin 27 1988-1989 N Wisconsin 31 Home Range Wolf / km2 1989-1990 N Wisconsin 34 1990-1991 N Wisconsin 40 Northern Minnesota 146 km2 24 1991-1992 N Wisconsin 45 Northern Minnesota 349 km2 33-111 1992-1993 N Wisconsin 40 Central Manitoba 253 km2 23-27 1993-1994 N Wisconsin 56 Northern Wisconsin 179 km2 41-82 1994-1995 N Wisconsin 83-86

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 337 away. A lonely wolf gives the “lonesome howl,” miles (8 km) per hour (Carbyn 1987). They a short call that rises in pitch and dies away usually trot or walk in nearly straight lines, with a sad note. If answered the wolf responds putting the hind foot in the track of the fore- with deep howling. Both alpha male and fe- paw. The winter territory size varies inversely male scent mark by lifting the hind leg. Other with the deer density (1980-1990) (see ac- pack members do not scent mark and urinate companying Fig.). While preying on deer the in squat positions. Two packs may merge for movement of wolves was 1-3 miles per day winter hunting. Pups normally do not join adults (Mech 1971a). The greatest movement was hunting until late September or early October, a straight-line distance of 886 km (531 mi) but spend the summer at outliers (= rendez- (Fritts 1983). Wolves move up to 45 miles vous sites) within the territory. (72 km) per day. Usually wolves move about Wydeven et al. (1995) report winter ter- 20 km per day (Mech et al. 1971). ritory size as 137 km2 on average (range 42- Estimates of wolf abundance can be made 287 km2) and a year round size 179 km2 by radio-telemetry studies, howling surveys, (range 49-323 km2). Mean dispersal distance and winter tracking (Wydeven et al., 1995). was 114 km (68 mi) for 16 wolves, the long- Counting wolves by “howling” techniques was est 480 km (288 mi), for a wolf that went pioneered by Bill Feeney and Clarence Searles north to Ontario. Fuller (1989) and Gehring in Wisconsin in April 1944 (Thiel, 1993). (1995) provide data on population dynamics Thiel and Welch (1981) began winter track- of wolves, mostly in Minnesota. ing to determine distribution, numbers, and A pack seldom exceeds a dozen individu- breeding status in Wisconsin packs. als and usually consists of fewer than that. In In Ontario the age structure was 35% recent years in Wisconsin mean annual pack pups, 40% yearlings, and 25% adults, where- size ranged from 2.6-6.2 (overall mean 3.9) as in Alaska it was 42% pups, 29% yearlings, (Wydeven et al., 1995). The density averaged and 30% adults (Mech, 1974). The survival rate about 19 wolves per 1000 km2 in occupied teris about 43% for pups, 55% for yearlings, and ritories. In 2000, there were 66 known packs. 80% annually for adults. Recent survival rates Wolves can run perhaps 35-44 miles per for Wisconsin wolves one year or older have hour, and for long distances at a rate of 5 averaged 82 % (Wydeven et al., 1995). Remarks. Although timber wolves occasionally prey on livestock and pets, so far the frequency of these occurrences is low. Fritts et Table Car-7. Wisconsin Home Ranges and Densities: Thiel 1993: 136 (In part based on Bill Feeney’s secret study). al. (1992) found a rate of 30 depredation complaints a year in Minnesota, for some 1200 Locality Years Range Size 1/x Wolf Density wolves and 7200 farms. In Wisconsin wolf Pleasant L 1940’s 186 29-93 depredation rates are generally less than one CampScott L 1940’s 225 32 percent per year (Wydeven, personal comm.). Willow Pack 1941-9 300 75 Wolves may at times contribute to the decline “ 1944-6 124 31-42 or suppression of deer populations, especially “ 1946-8 150 42-50 Michigan Co. 1941-50 90 45 in combination with severe winters and decline Twin Lakes 1944-46 86 — in habitat (Mech and Karns, 1977), but wolves Ave. 166 49 generally have limited impact on healthy deer Michigan populations. Because of its culling of the weak Cusino 1938 260 65 or sickly, predation at times may actually ben- “ 1950 75 25 Huldert 1950 48 8 efit deer. Tolerance or acceptance of wolves “ 1950 335 56 by humans only will be found in the remote Ave. 179 39 parts of Wisconsin. Wolves must be managed.

338 THE WILD MAMMALS OF WISCONSIN Additional Natural History. Thiel (1993) 1842. Vulpes fulvus: DeKay. The zoology of New reviewed the biology of the timber wolf in York. Mammalia, p. 44. Wisconsin, and Mech (1970) wrote a classic 1894. Vulpus pennsylvanicus Rhoads. Amer. nat., book on the wolf. 28:524. Not Vulpes pensylvanicus Boddaert, Geographic Variation. There is only one 1784, which is a gray fox (=Urocyon cinere- known geographic race in Wisconsin and oargenteus). His name is an invalid homonym Upper Michigan. and an invalid synonym. Also known as Vulpes Specimens examined. Total, 10. Dou- pennsylvania. glas Co.: Bear Lake, 2UW. T44N, R13W, 1758. Canis vulpes Linnaeus. Systema naturae, 1(1979). T44N, R12W. Sect. 20, 1 UW. 10th ed. T45N, R14W. Sect. 10, 1(1981)UW. Sum- 1960. Vulpes vulpes: Churcher. J. Mamm., 41:359. mit Twsp, T45N, R15W, Sect. 29, 1(1981) UW. Iron Co.: Paint Lake 1 UW. Oneida Co.: T37N, 1(1949). T36N, R8E, Sect. 12, 1 Vulpes vulpes fulva (Desmarest) (1949). McNaughton 1 (1930’s). See synonymy for the species. The larger fox of the Great Plains, Vulpes vulpes regalis Genus Vulpes Bowdich, 1821 Merriam, approaches the Mississippi River (Not Vulpes Oken) and its type is from Elk River, Sherborne Co., Swift and Red Foxes Minnesota. The name Vulpes means fox. The sub- “The Fox could not reach the grapes, although he specific name fulva means fulvous, an orange sprang again and again; and then, since he had not or ochraceous-chestnut color. the power, he left saying, The grapes are sour!” — Description. This medium-sized dog, Aesop’s Fable (about 600 B.C.), and the poet small for the several wild dogs in Wisconsin, Bahrius (200 A.D.) is best identified by color (see below) and long tail. The eyes have vertical pupils. The ears Medium to small wild dogs with long, pointed are long and pointed, tipped with black. All ears, tail round and bushy and about as long the feet are black. The front foot has a dew as the head and body. Color russet or rufous claw, and there are five toes. There are only red to sandy buff, with black feet and white four on each hind foot. The skull has a glob- underparts. Canine teeth elongate. The sagit- ular (subspherical) braincase with no sign of a tal ridges are V-shaped or nearly so. The type is Canis vulpes Linnaeus. Vulpes Oken is ruled invalid, as all his names are non-Linnaean.

Vulpes vulpes (Linnaeus) Red Fox

Concerning the fox hunt with horses and hounds: [Such hunting is] when people do the unspeakable to the inedible. — Oscar Wilde.

1820. Canis fulvus Desmarest. Mammalogie... pt. 1, p. 203, in Encyclopedia methodique... Type from Virginia. Skull of Vulpes vulpes.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 339 sagittal crest even in old adults (a mid sagittal ridge does develop). Differing from the similar skull of the raccoon (Procyon lotor), the snout of the red fox is slender and delicate, with elongate rostrum of the skull. There are slender but elongate canines. The rostrum of the red fox resembles that of the coyote, but is much smaller (see Measurements), and the domestic dog has relatively shorter canines. The gray fox Urocyon cinereoargenteus is similar in size, but instead of nearly V-shaped sagittal ridges they are lyre-shaped and prominently high (see that account). The red fox

Maps showing geographic distribution of Vulpes vulpes in Wisconsin and North America.

340 THE WILD MAMMALS OF WISCONSIN skull is not so flattened, and the rostrum is ceptionally bushy. Total length ranges up to narrower. There is no “step” on the ventral 1.5 m (58 in) and is usually 36-46 in. Ac- border of the dentary as in Urocyon. The tail cording to Schwartz and Schwartz (1981) is bushy. There are four pairs of mammae. body weights vary from 8-15 lbs (= 3.6-6.8 The baculum is troughlike, pointed at either kg). Ables (1975) reports 4.0-4.5 kg in fe- end. The chromosomes number 32 (Hoffmeis- males, and 4.5-5.4 kg in males. Males are ter 1989), or 34 (Rausch and Rausch, 1979).. larger than females. The normal color phase of the red fox is Dental Formula. DF = I 3/3, C 1/1, P bright rufous-ochraceous of the upper parts, 4/4, M 2/3 = 42. The incisors have minute black feet, and white venter and tip of tail. lobes distally. This tip is set off by scattered black hairs. Cross Geographic Distribution. The circumpo- foxes (with brownish fur and often having buff lar red fox may occur anywhere in natural patches near the legs, and a dorsal “cross” of areas, and in some human settlements. It vis- dark fur) and silver foxes (blackish frosted with its or dwells on islands in Lake Michigan and white-tipped guard hairs, and a white tail tip) Lake Superior. It occurs regularly and in wet- may differ from this pattern, and the former lands, highlands, forests, marshes, sand may somewhat resemble the gray fox. Espe- dunes, prairies, woodlots, forest edge. cially blackish silver foxes are often called Status. The red fox is managed by the “black”. Genes for the cross fox may be AA state as a game animal and furbearer (Pils and (red), BB (black), Ee or AaBBEe (Lariviere and Martin, 1978). Wisconsin and Michigan are Pasitschniak-Arts 1996). See table Car-8. Ju- in the regions where reportedly the great- veniles have dark gray-brown fur. Day length est fox harvests in the world are obtained. initiates the molt in April-June (Lloyd, 1980). It is an able predator of mice and rabbits, on In autumn the winter fur molts in, and the guard the obverse face of it, but kills chickens and hairs thin in March. Summer molt is loss of game birds on the reverse side of the coin. It the underfur giving the fox a ragged appear- is considered by many as the most glorious, ance in April-June. These phases are a single colorful, and beautiful mammal in the state. molt, with prime fur attained in winter. By those fortunate to observe the playful pups The red fox is about two to three feet at the natal den close at hand, the species is (600-1,000 mm ) in length. Although popu- an aesthetic blessing of nature. Its status is lar and technical literature is replete with state- excellent, due to proper management and its ments that the tail is half the total length, or own intelligence. From a sample of 203 red greater than the head and body length, I saw foxes from diverse areas of Wisconsin, Kube no evidence of that, even if one adds to the (1979) found a mean age of 1.6 years (with tail length the hair length at the tip. It is ex- some as old as 8 years), a replacement rate of 0.56, and that the species was replacing itself and increasing. However, Rolley (1993) reported that of 1,029 rural residents, responding to recent questionnaires, fox sightings recently had declined significantly (9 percent), which decline continued in 1994-1995 (Rolley 1996). In 1999-2000, the harvest was estimated from a survey of small game hunters as 11,614 (Dhuey, 2000) mostly from Wood, Shawano, and Brown counties. A trapper survey provided the estimate 7,717. The Skull of red fox, lateral aspect. Miller/Tertzi. increase in coyotes will probably cause red

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 341 fox populations to decrease somewhat (see cluding the opossums Didelphis, shrews Sor- Sargeant 1972; 1982). In American litera- ex and Blarina, and even Cryptotis (in Wis- ture the red fox is considered sly, wily, clever, consin—killed but not eaten), moles, cotton- and a chicken killer. tail rabbits, snowshoe hares, chipmunks, Like the coyote the red fox suffered from woodchucks, ground squirrels Spermophilus, bounty hunting. This practice began in 1923, tree squirrels, and flying squirrels Glaucomys, was repealed in 1931, reintroduced in 1959, beaver, mice Norway rats, muskrats, porcu- and repealed again in most counties in 1963. pines, weasels, mink, skunks, house cats, and Habitat. The diversity of habitats was carrion. Errington (1935) found mice and rab- detailed in the section on Distribution. Foxes bits in 25 Wisconsin stomachs. Pils and Mar- prefer a mix of woodland and meadow, and tin (1978) mentioned birds in their study of may thrive in a fragmented landscape. Today 132 stomachs and 1,020 scats. Birds preyed (Macdonald and Newdick, 1982) foxes are upon included herring gulls, grouse, ducks, found in some urban areas. Prey availability songbirds, poultry, and others. In southern is important for red foxes. For a comparison Illinois, Knable (1970) reported fox foods as of gray and red fox ecology, see account of eastern cottontails, (25% vol., 32% frequen- the gray fox (Follmann, 1973). cy); voles Pitymys (15%, 21 f.); Peromyscus Dens are usually found in remote, well- (6%. 15 f.), raccoons (6%, 10 f.); chickens drained areas, such as hay fields, hillsides, (9%); birds (15%); persimmons (10%); and shrubby fence rows, forest edge, and clear- grasses (16%). Red foxes eat 2.25 kg of prey ings. The natal den is usually a burrow used per week (Sargeant 1928). Food is eaten or for sleeping as well as rearing the young. A often cached for later use. fox family may share its burrow with another Reproduction. Arnold (1956) and Pils and fox family, even with badgers in England (Long Martin (1978) reported on the reproduction and Killingley, 1983). A typical den has at in red foxes in Michigan and Wisconsin. Storm least two openings. The burrow may extend et al. (1976) added information for the upper 30 feet, dug at a depth of four to 10 feet. Midwest. As many as 5-11 percent of females The nest is made of grasses in a widened part are barren (Samuel and Nelson 1982). They of the burrow. The burrow is often reworked are mature at 10 months but seldom breed from an abandoned burrow of a woodchuck the first year. Courtship begins as early as late or badger. A den has a conspicuous bare December. After courting by males, foxes pair mound of earth at the main entrance (Stanley up and, working together, dig a natal den. The 1963, Schmeltz and Whitaker 1977, Layne foxes breed from late December to mid-March. and McKeon 1956b, Pils and Martin 1978). Estrus lasts 1-6 days. There is a single litter Pils and Martin found five communal dens, per year. Gestation lasts 51-53 days. The lit- 11 percent of all dens found in southern Wis- ter is born in March or April (Sargeant et al. consin in 1972-75. 1981). Litter size is 4 or 5 pups, 5 or 6 (Pils Foods. An opportunistic predator and and Martin 1978), or up to an incredible high forager, the red fox primarily feeds on small of 17 pups (Holcomb 1965). (However, two mammals, reportedly even porcupines and litters from two mothers may share the same beaver (Payne and Finlay 1975), many birds, den and seem of one family.) Prenatal loss may even nesting herring gulls (Shugart and Scharf exceed 20 percent. If density is low, more of 1977), great horned owl (Pils and Klimstra, the red fox yearling females may mate. At birth 1975), and on carrion of large and small ver- the pups have their eyes closed, and the teeth tebrates. Insects, reptiles, and many fruits and hardly erupted. The tip of the tail is white. The other plant parts are eaten. Mammals used newborn weighs 71 to 119 g (Storm and Ables as food are summarized by Baker (1983), in- 1966). In ten days the eyes open, and at 20

342 THE WILD MAMMALS OF WISCONSIN days the young first emerge from the den to Automobiles kill many red foxes on the play like puppies. The fur is yellowish by 49 roads, perhaps including 25 percent juveniles. days. By 160 days the young are weaned but People often kill them accidentally with snow- brought small prey by the parents, remains of mobiles in Wisconsin. Farm machinery kills which are strewn around the den area. By some, mostly juveniles in dens. Many are shot October, young are full grown and have re- and trapped in the appropriate season. The placed the milk teeth with permanent teeth. regulated harvest usually is the chief mortality, They begin to hunt on their own and to dis- although locally mange may be the chief cause. perse from their place of origin. Males depart Home Range and Density. Home range first (Tullar and Berchielli 1980). The young for the red fox is large, especially for males. It can breed in their first year. varies from 142 to 1,280 acres. Winter spring Age variation in the red fox was report- home ranges obtained by radio-tracking for ed by Churcher (1960), Harris (1977), Pils 3 vixens averaged 1,495 acres. Foraging for and Martin (1987), and Storm et al. (1976). food leads to most wandering. The bound- Aging techniques and growth studies summa- aries of home ranges barely overlap. Home rized for Ontario show a huge proportion of range varies in Wisconsin from 57.5 to 161.9 juveniles (Voigt, 1987). Kube (1979) discussed ha for 7 foxes radio collared. An adult male age classes for Wisconsin (see Status above). used 5.12 km2. Home ranges are usually twice Mortality. Man is the worst enemy of the as long as wide (Follman 1973, Harris 1977). red fox (Pils and Martin 1978). Occasionally, Ables (1969) found home range significantly but rarely, large predators such as coyotes and smaller in urban areas. On an average night wolves may kill one. Badgers prey on the a fox on snow may travel 5 miles (= 8 km) pups. Raptors may do the same. A real dan- (Arnold and Schofield 1956); the winter home ger for Vulpes vulpes is sarcoptic mange (Sar- range was 900 acres (= 364 ha). coptes scabei). In winter the denuded foxes Territories (approximate home ranges in die of cold. On Washington Island hunters this example) are defended by scent marking, reported mange infecting red foxes in the not by combat. Follmann (1973) found defend- winter of 1992-1993, and a decline in fox ed territories to average larger in males, and numbers resulted. Pils and Martin (1978) to be of greatest size in January (1,091 acres found only 7 percent infected in Wisconsin, = 2,357 ha). The mean annual size was 626 but the carcasses are hard to find. Another acres (253 ha), with five months missing data. potential danger is rabies. It had a great im- For females the annual size was 278 acres (112 pact in Europe in the 1970’s. An occasional ha). The range was 172 to 361 acres for fe- fox is rabid in Wisconsin. Canine distemper, males, and 290 to 1,091, in males (see Table parvovirus, canine hepatitis, toxoplasmosis, Car-9). Caches are marked with urine scent tularemia, and encephalitis kill foxes (Voigt, (Henry, 1977). Territorial boundaries are vis- 1987). Coyote diseases may be transmitted ited regularly, at about 2-week intervals (Mac- to red foxes (Sargeant, 1972, 1982). Donald, 1979). Homing is commonplace in Parasites include mites, ticks, sucking lice, displaced foxes. Although red foxes and coy- six fleas, tapeworm, heartworm (Dirofilaria otes forage on the same ground, the foxes immitis), Paragonimus, numerous flukes, avoid contact (Voigt and Earle, 1983). roundworms, and a flagellate protozoan Annual reproduction has been estimat- (Erickson 1944, Goble and Cook 1942, Mor- ed at 214 percent per adult (Layne and McKe- gan 1944, Haas 1970, Scharf and Stewart on 1956). Mortality of young is high, and it 1980, Jackson 1961, Baker 1983). Ascarids continues in adults, so that young make up and hookworm Uncinaria affect many pups most (74 percent) of the fur harvest (Pils et (Voigt, 1987). al. 1981). Foxes may live in nature between

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 343 three and eight years, reportedly five years Table Car-8. Two factor inheritance in red foxes. All color (Storm et al., 1976; Lariviere and Pasitch- phases can occur in the same litter. Silvers are found on fur niak-Arts, 1996). There seems to be a peri- ranches. Ashbrook, 1937, and Lariviere and P.-Arts, 1996. odical increase in foxes approximately every Ashbrook: 10 years (Arnold 1956). Mange may cause Red AA BB sudden decreases of fox numbers in the cold AA Bb (smoky red) winters. In southern Ontario, when mortality Cross Aa BB Aa Bb is high, more red fox yearlings breed (Voigt Silver AA bb and MacDonald, 1984). aa BB Rolley (1993) reported the results of Aa bb about 1,000 questionnaires sent to Wiscon- sin farmers. The average number of litters aa Bb aabb statewide since 1986 holds to about 1.5 families per farm. Lariviere & P.-A: Arnold (1956) describes the autumn move- Red AA Cross AABB, Ee, AaBBEe ments as the “fall shuffle.” The young disperse Black BB in late September and early October from the dens, males a month before females. Males may move an average 18.4 mi (29.4 km) from the ture Sanctuary at Jackson Harbor, all sight den, and females average 6.2 miles (9.98 km). records), Florence, Iowa, Jackson, La Some juveniles move as far as 126 miles (202 Crosse, Manitowoc, Marathon, Marquette, km) (Longley 1962) and reportedly 245 miles Monroe, Oconto, Oneida, Portage, Price, (392 km) from Wisconsin to Indiana (Ables Racine, Sheboygan, Trempealeau, Vernon, 1965). (Not only is this distance amazing, but Washburn, Washington, Waupaca, Wood how did the fox avoid Chicago?) counties. Family units often vary from six to ten Other Records (Balliett and Taft, 1978): individuals. Occasionally related families may Fond du Lac, Oneida, Winnebago and Lang- form a merged group. Pils and Martin (1978) lade counties (not plotted). estimate 7 red foxes per 1,000 hectares. Michigan. Schoolcraft Co.: 4 mi. N Remarks. Even paired red foxes hunt as Manistique 1. Big Summer Island in Lake individuals. The hunting is nocturnal, although Michigan, 1. occasionally they hunt in daylight. Usually the fox emerges from its resting burrow at twilight. When the fox locates prey often it leaps Genus Urocyon Baird high in the air to pounce on it. Gray Foxes Additional Natural History. Pils and Mar- tin (1978) reviewed the natural history for the “A gray fox... stomach content: a whole ground red fox. Lariviere and Pasitchniak-Arts (1996) squirrel well chewed plus one lizard foot and provide a recent review. somewhere inside the ground squirrel a bit of Geographic Variation. There is a single aluminum foil. The secret.”— Gary Snyder. Turtle subspecies of red fox in Wisconsin and Upper Island, 1974. Michigan, but a larger red fox ranges up to the Mississippi River from the western plains. Baird’s description (1858) refers to the bris- Specimens examined. Total, 43. Adams, tly line in the tail, dental peculiarities, and the Ashland, Barron, Chippewa, Columbia, step in the dentary. These may differ only in Dane, Door (Washington Island, at Airport; degree from Vulpes (Seton, 1953), but the at Pedant’s Lane; at E. Harbor; Ridges Na- genus seems distinct.

344 THE WILD MAMMALS OF WISCONSIN Urocyon cinereoargenteus (Schreber) gray fox is unique in North American Canidae Gray Fox in two characteristics, the U-shaped or lyrate sagittal ridges and an extra step or notch in Urocyon cinereoargenteus the dentary in ventral outline. Some other fox- cinereoargenteus (Schreber) es approach these conditions. A good character is the short muzzle, with short nasal 1775. Canis cinereo argenteus Schreber. Die bones and delicate antemolars. Concerning Saugthiere..., Theil 2 (Heft 13), pl. 92. Type the lyre-shaped sagittal ridges, there is more locality vague: Eastern North America. than merely a curved outline (compared to 1775. Urocyon virginianus Schreber. Die Saugthiere... the typical V-shape in Vulpes vulpes), for the Theil. 3, p.361, 585. Type from Carolina... ridges steeply protrude from the cranium on 1894. Urocyon cinereo-argenteus: Rhoads. Amer. their inner and outer margins, and below the Nat., 2l8:524. Combining adjectives into one ridges (ventrolaterally) the bone is textured name. with a pebbly character. There are reports of 1899. Urocyon cinereoargenteus ocythous Bangs. black phases of Urocyon, and Root (1981) Proc. New England Zool. Club, 1:43. Type reported a male, young-of-the-year Sampson from Platteville, Grant Co., Wisconsin. Synon- gray fox (lacking guard hairs) from Richland ymized here. County, in 1979. The chromosome number differs from The name Urocyon means dog with a Vulpes vulpes, in diploid number (66) ac- tail, but of course all dogs have tails. The ques- cording to Fritzell and Haroldson (1982). The tion is, what sort of tail? Likely the name re- baculum is dugout shaped and shorter than fers to the dark, bristly line on the dorsal sur- in Vulpes. face. It may or may not be tipped with black. The color is distinctive, grizzled gray The long name cinereoargenteus means gray above, ochraceous orange laterally, and whit- and silver, referring to the dorsal fur. ish below. The dorsal underfur is rather buffy Description. The gray fox is a small, stout and the guard hairs are also buffy basally, but canid comparable to the red fox Vulpes, but blackish outward, becoming white distally. In differing in grizzled, grayish color, rounded, many specimens the tips are black. Therefore, shorter ears less defended by guard hairs, and there is a mixture of black and white, and the muzzle shorter, broader, with smaller teeth. overall tone of gray. There is black around the The legs are shorter, and the gray fox is slight- eyes, usually encircling the muzzle, and the lips ly smaller than the red fox. The skull of the are black. On the tail the dorsal hairs are so

Photo of gray fox by Allen M. Pearson. Skull of Urocyon cinereoargenteus.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 345 stiff and so black that a long band extends to southern half of the state (except for rare oc- the tip of the tail, which is usually blackish (not currences) in the 1970’s. Mappings from that white as in Vulpes). The feet are dark brown- time were based on harvest and observations ish or blackish but not as black as in Vulpes. (Petersen et al., 1977). Jackson (1961) be- The nosepad is black. Tawny orange extends lieved that the grays had invaded the north- along the sides and from the ears ventrally, ern counties, with rare occurrences, from encircling the white chin and throat, compos- 1910 to 1960. ing a handsome collar of orange. The ventral In the far northern counties Florence and fur is white. There is a single molt with prime Forest, McCabe (1972) mentioned that the fur attained in winter. first DNR records, beginning in 1927 show a Gray foxes usually weigh 9-11 pounds few grays and no red foxes. Eleven years lat- (4.1-5 kg) and are about a meter (39 in) in er red fox harvests increased “markedly” as total length. The tail is about 15 inches. Males the gray fox “declined”. This trend continued are only slightly larger than females. An adult until at least 1972 (see Fig.). Root (1981) sug- male from Fort McCoy measured in total gested grays in southwest Wisconsin were in length 1160, tail length 440, length of hind decline, and advocated a restricted harvest. foot 155, and ear length 62 mm ( = 46, 14, Gray fox pelts were priced high at that time. 7, 2.4 in.). This male and two adult females Pelts are coarse, usually worth less than those from Portage and Washington counties had of red foxes. cranial measurements as follows: condylobasal In the 1970’s, Petersen et al. (1977) length 130.3, 126.4, 121.5; length of nasals found gray fox populations throughout Wis- 50.7, 39.9, 44.9; zygomatic breadth 72, consin were stable or declining. The gray fox 71.3, 68.4; interorbital breadth 26.4, 26.7, then showed its highest abundance in the hilly 24.5; maxillary toothrow 55.1, 53.6, 51.2. southwest part of the state, and in the Kettle Dental formula. The gray fox has a typi- Moraine and adjacent agricultural areas of cal number of teeth for Wisconsin canids, DF southeast Wisconsin. Gray foxes were rare, = I 3/3, C 1/1, P 4/4, M 2/3 = 42. by observations or harvest records, in north- Distribution. The gray fox occurs through- ern counties. In central Wisconsin they were out Wisconsin, ranging northward onto the present (Long, 1970) and not considered Upper Peninsula of Michigan. The primary scarce by a local fur buyer. See status map by range is southern and central Wisconsin. In Petersen et al. 1991, the gray fox established itself on Wash- Today the general distribution is about ington Island, probably crossing lake ice from the same, with highest abundance in the Upper Michigan or the Door Peninsula. southwest counties and fair numbers in the Status. Although the gray fox is known Kettle Moraine area. The species ranks sixth from throughout most of Wisconsin, and among harvested furbearers (behind muskrat, much of the Upper Peninsula of Michigan, it beaver, raccoon, red fox, and coyote). Ap- is rare in northern counties. In several studies parently the number of gray foxes in Wiscon- on the status of the gray fox, reviewed by sin has steadily increased with protection and Petersen et al. (1977), the gray fox seemed management. In Wisconsin, there is an esti- practically confined to the Mississippi and mate of 12,400 grays (Petersen, personal Wisconsin river basins (up to 1921), with some corr.), and the harvest goal is about 5,100 scarce reports later from central, eastern, and animals Crossley, 1985). In 1975, Petersen Door counties (1953). These mapped extral- et al. estimated that only 3,000-4,000 foxes imital records were based on subjective opin- occurred in Wisconsin. Dhuey (1994) reports ions of wildlife managers and in bounty a recent annual harvest by trappers (1,251) records. The primary range contracted to the and hunters (475) of 1,726 gray foxes. In

346 THE WILD MAMMALS OF WISCONSIN 1999-2000, the harvest was estimated as 1,553 (Dhuey, 2000), continuing the downward trend. Although pelt price relates to the harvest of gray foxes, another direct relation is seen in the demand for red fox and raccoon furs (Petersen et al, 1977). Fritzell (1987) suggests gray foxes may survive with harvest of 55 percent, but usually less than that would be surer. The status of the gray fox seems good, although Rolley (1996) found evidence that foxes recently declined. Although human centers such as Milwau- kee, Madison, Appleton, and even smaller

Maps showing geographic distribution of Urocyon cinereoargenteus in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 347 cities probably eliminate habitat for gray fox- the dominant fox on the island, according to es, this adaptable species persists in protect- numerous residents. I have seen 1-9 observa- ed adjacent areas, such as the Kettle Moraine tions of grays each summer since 2002(one State Forest and nearby farmlands. Fritzell dens near my house presently on the north (1987) discusses the interesting biology of this shore) and not a single red fox in that time fox, adapting to land use such as farming. (except a road-killed red fox in June 2002). Especially in southern and central Wisconsin, In Follmann’s (1973) study in southern famous for dairy farms, there is abundance of Illinois, red and gray foxes were sympatric, food (cottontails, mice, corn, and apples) and many groves of trees, brushy areas, and hedgerows providing cover. These seemed to benefit gray foxes. In the rugged bluffs and hills of southwest Wisconsin conditions are ideal for this species. Habitat. The gray fox is a woodland canid, agile enough to climb trees (Jackson, 1961). Nevertheless, the original range seemed restricted considerably from northern forests. In some situations gray foxes may outnumber red foxes (Follman 1973; Tullar and Berchielli 1982). The author has seen little evidence of that in Wisconsin, where the red fox harvest may dramatically increase while the gray fox numbers fall. For a comparison of the red and gray foxes, Follman (1973) presents a careful study. Smaller than the red fox, and less adaptable to human activities, the grays are less common in Wis- consin, seldom seen, and seldom killed by Graph Showing Gray Fox Abundance 1975-1976. hunters and automobiles. They persist in num- Petersen et al. Recent status changes – see text. bers cycling irregularly in upland hardwoods and riparian mixed hardwoods in southern counties, inhabiting hardwoods, cedar swamps, tamarack stands, and mixed coniferous forests in northern Wisconsin, and on the Upper Peninsula of Michigan (Jackson 1961, Wenzel 1912). The gray fox thrives in brushy areas near woodlands. Its preference for rough (rocky) terrain may be another reason (the big-timbered bottomlands of several rivers is likely another) that so many occur in southwestern Wisconsin. On Washington Island, where red foxes were common and doubtless cross the ice from one isle to another, the grays surprisingly showed up in 1990-1991. Since then, Fox harvests in Florence and Forest counties. McCabe until August 2002, the gray fox seems to be 1972. A competition not well known.

348 THE WILD MAMMALS OF WISCONSIN and the reds usually (except in March, the time pines (Weaver, 1939) and eat shrews. They of breeding vixens) had larger home ranges occasionally capture birds, amphibians, wea- or territories. The grays were expected to re- sels, and numerous rodents (Errington, 1935). place the reds eventually because they have In season plant foods such as elderberries, more omnivorous diet and use a greater vari- grapes, acorns, grain, and apples are eatern. ety of dens. In winter the grays kept to the Pils and Klimstra (1975) found in autumn that oak-hickory highlands most of the day (54 70 percent of the diet may be vegetation. percent use, and especially in daylight), while Reproduction. The mating season of the the reds also used old field and woods during gray fox lasts from late winter until March, the day. In spring the grays ranged into the and gestation perhaps lasts about 53 days fields at night to forage (47% woods to 26% (Layne and McKeon, 1956; Richards and old fields). In summer both reds and grays were Hine 1953; Sheldon 1949). Fritzell (1987) usually in the old fields at mid-day. In autumn reports 59 days, based on captive foxes. The the use of habitat was variable, but grays were den is prepared by male and female and she in the upland woods at mid-day (47% use) bears on average four pups (1-7). Richards when reds were there the least. and Hine (1953) determined the litter size to There seems little information on gray be 3.9 for 44 females, and Root (1991) found fox dens, which are occasionally excavated 3.5 for 85 females. (Both studies were of pla- near a stream in woodland (Hoffmeister, cental scars, which is a little higher than true 1989). Dens are often appropriated from litter size.) About one fifth of the yearling fe- another mammal, and seldom if ever found males do not breed (Root, 1991). In other in fallow fields, cultivated fields, prairies, or studies about 8 percent do not breed (Fritzell, meadows. Often gray foxes den in hollow 1987). The pups are nearly naked at birth (in stumps and logs, rock crevices, even brush May), their eyes unopened. They begin to and woodpiles. I do not know if they line their grow a wooly juvenile pelage. From about 100 nests with leaves and grasses as the red fox grams, each pup grows rapidly, and after three does, but it seems likely that they would. weeks prey is brought to them by the parents Crossley (1985) reports that dens are typical- to eat. Weaning takes place in about 80 days. ly lined with grass, leaves, or shredded bark. The baby teeth are replaced by four months, Occasionally the gray fox will rest in the day- and then the offspring forage alone. Males time in dense brushy cover. Most dens found first, and for longer distances, disperse in au- in southwest Wisconsin are sited on the east, tumn, followed by young-of-the-year females. south, or southeastern slopes of the hill (Frit- Longevity is 10-11 years (Crandall, 1964), zell, 1987). The gray fox often sleeps during as in red foxes. winter days in excavated dens. The den may Mortality. Humankind, of course, traps be used year-round. gray foxes for fur, hunts them, and occasion- Foods. Gray foxes are somewhat omniv- ally kills one with an automobile. As much as orous. They are not likely to raid a chicken half the population of gray foxes in some lo- yard, a habit which has made the red fox so calities of Wisconsin has been harvested in a despised. They are less likely to feed on game single year. Any large meat-eater that might birds and waterfowl. Their primary foods are prey on the red fox likely would prey on the mice, cottontail rabbits, fruits, mast, and ar- gray, especially the pups. Pups show the high- thropods (especially insects, grubs in winter) est mortality, over 40 percent. Numerous and considerable carrion (Errington, 1935; parasites are known from the gray fox, sum- Hatfield, 1939; Richards and Hine, 1953). marized by Jackson (1961) and Baker (1983). In winter the chief food is the cottontail (Cross- These include lice, fleas, ticks, and mites. In- ley, 1985). Reportedly grays attack porcu- ternal parasites include flukes, tapeworms,

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 349 roundworms, spiny-headed worm, and occa- to be declining there. Root (1981) found gray sionally heart worms. Some gray foxes are foxes in southwest Wisconsin in decline, re- infected with rabies, and numerous other dis- portedly from high trapping mortality. He eases such as plague, tularemia, St. Louis (1981) calculated a composite life table for encephalitis, Q fever, listeriasis, leptospiro- the gray foxes in southwestern Wisconsin for sis, histoplasmosis, toxoplasmosis, canine November-January 1978-1980. Of 636 ani- hepatitis, and Tyzzer’s disease. Infections can mals, 420 were young of the year, and 118 be a major cause of mortality, but gray foxes were yearlings. Thus, two thirds were young- are quite resistant to sarcoptic mange. They of-the-year and 85 percent comprised the first seem to suffer little infection from heartworm two age classes. Trappers collected 243 gray (Fritzell, 1987). foxes in southwestern Wisconsin. The calcu- Home Range and Density. Richards and lated total mortality in a year was a surprising Hine (1953) found home range varying greatly 66 percent. in Wisconsin, from 145 acres to as much as Additional Natural History. Fritzell and 2,155 acres (59 to 872 ha). During winter Haroldson (1982) reviewed the gray fox. courtship wandering is more extensive, and Geographic variation. There is none ev- around the natal dens the wandering is less. ident in Wisconsin. The question whether Jackson (1961) and Crossley (1985) give the there is a race U. c. ocythous differing from Wisconsin home range size as 31-765 acres. the nominate race found in Illinois was raised In southern Illinois, Follmann (1973 by Hoffmeister (1989) who referred all foxes found territories larger in males, and greatest he examined to the nominate race. No one size in April (to 501 acres). The annual size has addressed the problem in nearby Nebras- of territory was 336 acres for males, 265 for ka, Iowa, Minnesota, Missouri, Wisconsin, or females. The range was 178 acres to 414 Michigan. Cory (1912), Jackson (1961), and acres for a female, and for males 177-501 Long (1974) referred the Wisconsin speci- acres (see Table Car-9). mens to U. c. ocythous mostly on geograph- The wild population probably consists ic grounds. Burt (1946) surely relied on geo- chiefly of young-of-the-year (Lord, 1961b). graphic grounds, because the rare grays in Some foxes may live six years, possibly sev- Upper Michigan doubtless emigrated from en. The populations cycle multi-annually, Wisconsin. One wonders just how far the con- some peaking approximately every 10 years. cept of geographic grounds has served to map In Wisconsin there is no evidence of such cy- this race (Hall, 1981), from southern Canada cles. Populations seem usually low, for the southward into Oklahoma and Arkansas. State animal is seldom observed. Males have larger boundaries seem influential in fixing racial home ranges, and wander less than Vulpes boundaries. vulpes. McCabe (1972) found in the years The Upper Mississippi Valley gray fox, 1937-1955, in northeastern Wisconsin, near with type at Platteville, Wisconsin, was de- the Pine and Popple rivers (in Florence and scribed as pale. Ostensibly the pale foxes Forest counties), that gray foxes were in de- range into the southern Great Plains as far as cline, and red foxes, judging from trappers’ Oklahoma. A specimen from Oklahoma in reports, were concomitantly increasing. this collection was decidedly pale, both in the Whether competition is indicated is unknown, whitish hairs of the dorsal fur and the pale for heavy cutting of timber in the region might ochraceous (rather than rufous) of the throat cause the same effect. I have observed gray and sides. The race U. c. ocythyous also is and red foxes together on Washington Island supposed to have long tail, long ears, and slen- from the 1990’s until the present, and the der muzzle. These are trivial, especially in gray foxes are thriving. Indeed, red foxes seem carnivores, where such differences are usual-

350 THE WILD MAMMALS OF WISCONSIN ly slight and the character highly variable. In Clark, Dane, Door (Washington Island), Hall (1989) the race U. c. ocythous is mapped Dunn, Jefferson, Juneau, Monroe, Pepin, from Platteville, Wisconsin (the type locality) Portage, Washington counties. through Wisconsin and most of Upper Mich- Other records. Jackson 1961; Long, igan, into northwest Illinois, across the Plains 1974; Root 1981 (no voucher specimens to eastern Wyoming, north to Ontario, and preserved). south to the Texas border. The specimens I have seen from Wisconsin are not pale, all having the same bright coloration. Although Family URSIDAE Gray 1825 assigning Wisconsin foxes to the pale race, Bears Jackson (1961) examined the holotype in the Museum of Comparative Zoology at Harvard, The bears are large, short-tailed carnivores and reported that it was “greatly faded”. The with robust canines and bunodont cheek teeth. faded specimen was thought to represent pale The skull otherwise resembles those of large, Great Plains foxes. In the USNM, I noted pale stout dogs or wolves. gray foxes from Oklahoma and the Great Plains, but Wisconsin foxes were darker as are my specimens. The Mississippi River may Ursus americanus Pallas be roughly the border for the pale foxes, or Black Bear the racial boundary may lie even westward or southward. All Wisconsin foxes are referable “The black bear (U. americanus, Gm.) of North to U. c. cinereoargenteus. The name U. c. America is well distinguished with a flat forehead, ocythous is a synonym, and the name of the smooth and black fur and fulvous muzzle.” — Baron pale foxes on the Great Plains must be Georges Cuvier, in Regne animal, 1846. changed to something else. They seem referable to the southern U. c. scotti (western 1780. Ursus americanus Pallas. Spicilegia zoologi- Texas), which is a pale, prairie race. ca... Fasc. 14:5. Type from eastern North Specimens examined. Total, 17. Adams, America. Chippewa (Cadot, specimen not preserved). 1961. Euarctos americanus americanus (Pallas): Jackson. Mammals of Wisconsin, Univ. Wis- consin Press, p. 311.

Table Car-9. Mean territories [i.e., home ranges] of foxes in acres. Follmann 1973. Ursus americanus americanus Pallas Month Gray foxes Red foxes 2 males 4 females 2 males 6 females The name Ursus means bear, a Linnaean Jan 202 262 1091 321 appellation, and americanus of course means Feb 488 368 838 229 Mar 485 414 631 208 Apr 501 179 385 392 May 452 228 417 311 June 253 297 733 328 July 259 — — 330 Aug 380 — — 361 Sept 177 — — 296 Oct — 178 — 177 Nov 243 204 — 172 Dec 256 255 290 216 Yearly 336 265 626 278 Black bear’s bunodont cheek teeth. S. F. Baird, 1858.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 351 tem of dentition, almost frugivorous, explains why the bears seldom devour flesh. Students often confuse the skull of a young bear with the massive skull of a male wolf, but one look at the carnassial upper premolar will tell the story. The bear’s cheek teeth are all broad with low cusps for crushing, and they are not developed at all for shearing meat. The rostrum is more robust than that of the wolf. The anterior premolars are separated by gaps, tending to be small, vestigial; Black Bear by Georges Cuvier, Regne Animal, 1846. and often one, two or more are lacking. Cu- Woodcut. vier noted that the cartilage of the nose is elongated and moveable. There are three pairs an American origin. The word bear is derived of mammae, two pairs pectoral and one pair from Old English brun, or brown. Black does inguinal. The last upper molar is less than an not always apply; many bears are brownish inch (25 mm) in length in the black bear (it is or cinnamon, and often are spotted on the greater than an inch in the grizzly). The bac- throat or chest with white ulum is not especially large, is rather club-like Description. The huge size of the black (Burt , 1980), and tapers to either end. Slightly bear and its well-known stout, “roly poly” form curved, it measures 140-167 mm in length. make identification easy. When the adult black The fur is deep, glossy black with patches bear stands on its hind feet and lifts its mas- of white ranging from a small spot to a large sive head high, it seems less “roly-poly”. It has a short tail, long claws on all four feet, and dense coarse fur. The ears are rounded. The massive feet are broad, naked-soled, pentadactyl, and plantigrade. The foreclaws are recurved. The eyes are relatively small, and the rhinarium or nosepad is broad. The skull is correspondingly large in adults, wolf-sized in subadult bears, and usually the largest of any Wisconsin carnivore, in length, width, and height. The teeth are quite distinctive, because the cheek teeth are bunodont in relation to the varied diet (see below). The canine tooth exceeds 15 mm in length. Long ago in Regne animal, the great anatomist Baron Georges Cuvier (1846) provides this thorough description of the dentition: “The Bears (Ursus, Lin.)—Possess three large molars on each side of both jaws, altogether tuberculous, and of which the posteri- or above are most extended. These are pre- ceded by a tooth a little more trenchant, which is the carnivorous tooth... and by a variable number of very small false molars... This sys- Skull of Ursus americanus.

352 THE WILD MAMMALS OF WISCONSIN blaze found on the chest of about half of the Wisconsin bears. The muzzle is often cinnamon brown instead of black, and there are many bears that are cinnamon brown, deep walnut brown, or even a yellowish dark brown. Molt occurs in April or May, and the new fur is acquired by June. The hair continues to grow out until prime in October, when it is not only longer but darker in color. In the spring the prime fur soon wears, fades, and falls away. Adult males weigh about 250-350 pounds (113.4-159 kg), and females 120-180

Maps showing geographic distribution of Ursus americanus in Wisconsin and North America. The range area has shrunk.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 353 pounds (54.5-82 kg). Large males rarely ex- sons for hunting of bears with guns began in ceed 600 pounds (273 kg). Total length rang- 1934, and for bow hunters in 1942. Spring es up to 1,780 mm, nearly six feet in total trapping was allowed from 1945-1956. Hunt- length, but usually bears are smaller. ing and trapping seasons protect mammals Dental Formula. I 3/3, C 1/1, P 2-4/2-4, at least for much of the year. In 1957, bear M 2/3= 34-42, with the full complement 42. trapping was entirely prohibited, and in 1965, Geographic Distribution. The black bear cubs were protected. In 1985, the bear sea- formerly occurred throughout the state and son was closed, although some poaching was the Upper Peninsula of Michigan. It disap- reported. Since then the harvest has been peared from southeastern Wisconsin by 1860, closely regulated and the bear population has as a result of human activities, and from cen- more than doubled. Mothers with cubs are tral and western Wisconsin by 1890. The now protected. Hunting and number of bears statewide bear population probably reached harvested are regulated in management zones. its lowest point around 1915. Since 1950, There are two separate seasons of taking black bears have increased to thousands in bears, one with the help of dog packs and northern Wisconsin, and are expanding their the other using bait or tracking, with no dogs. geographic range southward, where bear- Chippewas were granted the right of tribal human interactions will become problems. In hunting of bears since 1983. In Upper Mich- 1961, Jackson determined Wisconsin’s pri- igan, 37 percent of the bears taken were killed mary bear range to be bounded by a line from in their winter dens. Polk County down to southern Clark Coun- An injured bear or a mother with cubs ty, eastward through Marathon County, and can endanger human life, and should not be on to Green Bay near Marinette. There was approached. Encounters occur often because one report from the Apostle Islands in Lake a bear smells food. Bears can easily outrun a Superior, and none on Washington and Rock human. Black bears can readily climb trees; islands. By then bears had vanished from Door therefore, climbing is no escape. Fortunate- County. Some transplants were made in Door, ly, a bear usually runs away when it encoun- Adams, Jackson and Wood counties (Jack- ters a human. An adult who encounters a bear son, 1961), but the revival of bears is best usually should act unconcerned, neither ag- attributed to game laws and increased human gressive nor afraid. A child should scream acceptance. In the past quarter of a century, defiantly, hopefully to bring help. In wilder- the range of Ursus has expanded southward ness, children should always be attended by into Wisconsin’s central forest lands, and bears adults. Jackson (1961) and Baker (1983) occur occasionally in nearly every non-urban mentioned a three-year old Michigan girl killed county in the state. In the past decade three and partially consumed by bears. The Milwau- bears have swum over to Washington Island, kee Journal-Sentinel (Aug 21, 2002) discuss- and two of them were shot dead. One was es a baby killed out of state, and mentions seen swimming between Plum Island and the that in Walworth County, Wisconsin (August Mainland (see Fig.). They also occur on some 1999), a boyscout was dragged 80 feet with Apostles, especially on Stockton Island. his tent, bitten and badly injured (dislocated Status. See comments in Geographic shoulder). Distribution above. Schorger (1949) reviewed Bears occasionally damage beehives, tear the history of black bears in Wisconsin, and up property such as barn doors, and kill live- W. E. Scott (1947) discussed their status about stock. Pigs, sheep, cattle, dogs, and horses that time. Briefly, bears were protected in might be killed by a hungry bear. One tried to Wisconsin by a hunting season in 1917-1918, enter a house through a window at Lake DuBay but usually unprotected prior to 1930. Sea- and was shot. In Wisconsin a farmer may be

354 THE WILD MAMMALS OF WISCONSIN compensated up to $5,000 for bear damage Home Range and Density below. Briefly, under the Wisconsin Wildlife Abatement and bears are considered a threat by rural land- Damage program. Approximately 1,000 nui- owners, farmers, and bee-keepers, as a mag- sance bears or more in Wisconsin have been nificent game animal by hunters, and as a rare, moved to remote locations in the years 1996- but popular, wild mammal by many natural- 1998, following complaints to the Department ists. In my opinion, this species is well man- of Natural Resources. The transplants were aged in Wisconsin, which does not want a carried out by the U. S. Department of Agri- surplus of bears near people. culture, Animal Damage Control. Bears can Habitat. The black bear prefers heavily be bothersome in numbers around garbage forested areas, hardwood and mixed hard- dumps, and the ethics of shooting garbage woodforests, dense brushland, rocky terrain, dump bears often has been debated in Wis- and swamps and occasionally is found in re- consin. Presently it is illegal to hunt at dumps. mote marshes. Bears change from one habi- The bear hide is seldom worth as much tat to another in search of berries in season as $100, and the hair is so coarse the pelt is and other plants (Rogers, 1987). Forest open- usually made into a rug or wall hanging. The ings, then, are useful as a source of berry fur historically has been used for coats, trim, foods. Oak stands provide hard mast. hats, and lap robes. The meat is excellent, The bear in summer usually finds a hid- unless tough and stringy from an old bear, or ing place for rest, its lair, where it occasional- too fat. The hams resemble beef steak. Bear ly scratches the earth or piles vegetation for a fat or grease was formerly used as a salve to bed. When food becomes scarce in late fall a prevent mosquitoes from biting, as a cure for bear, usually obese with autumn fat, prepares arthritis, and to preserve leather, and the lard a den for its winter sleep. This hiding place is was useful in cooking. usually insufficient in size to accommodate the Some statistics on the harvest of Wiscon- bear’s entire body although if lucky it may find sin bears are given in Tables Car-10-11, from a cave or hollow tree. Usually a partially ex- Dhuey and Wallenfang (1995) and Kohn et cavated pit at the base of a wind-blown up- al. (1998). In 1999, the harvest was reported rooted tree or even dense vegetation such as as 2,881 bears (Kohn and Rolley, 2000). See evergreens provides what shelter there is, and the bear is eventually partially covered with snow. I have a photo of this kind of “den”. A DNR warden (winter 2004) reported a Wis- consin bear denning in an eagle nest, and when the eagles returned to it the bear had not wakened. Kessler (1994) reports 62 percent of dens near trees or windfalls, 25 percent excavated, and only 8 percent unprotected as “nests.” The pregnant female deliv- ers her litter of cubs in the winter den, which, therefore, is also a natal den. Foods. Jackson (1961) lists a variety of foods including small mammals, bird eggs and broods, garbage and carrion, berries of all kinds, nuts such as beech nuts, acorns, and hazel nuts, ants and their young, honey, fishes, and many other things. Twigs, berries, Abundance of black bear in Wisconsin. After Wise. grapes and other plants, and rarely fungi, are

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 355 eaten. Other foods are earthworms, insects, confirms plant foods such as grasses, sedges, mice, nests of hymenopterans, fish caught serviceberry, wild sarsaparilla, blackberry, and with the bear’s mouth, frogs and salamanders, raspberry. Ants and even deer (13 percent of turtles and their eggs, hares and rabbits, wood- scats) including two fawns were eaten. chucks and other squirrels, snakes, other bears Reproduction. Bears breed promiscuous- rarely, deer (fawns and some adults). Altogeth- ly in summer (June and early July), mating er the vegetable matter may be 77 percent, and pairing temporarily. Females may pro- carrion 15 percent, insects 7.4 percent, and duce litters, depending on sufficient food avail- small mammals almost 1 percent (Banfield ability, by three years, usually four, and as 1974). Some bears feed on livestock. In the long as seven (Kolenosky and Strathearn, Apostle Islands they have been reported to 1987). Females with young avoid adult males. raid beaver lodges (Smith et al., 1994). Nursing females do not conceive, therefore Baker (1983) discusses the foods of black females breed in alternate years. Bears be- bears, mostly from studies in places other than come sexually mature at three years to three Michigan, saying the black bear has a reputa- and a half years of age. The dominant male tion for being continuously and ravenously fiercely drives off competitors. Bears are nev- hungry. Blueberries and acorns were of such er monogamous for long periods. Implanta- importance to influence, by scarcity, depre- tion is delayed 4 1/2 months with young born dations on livestock and gardens by hungry January 15-February 15, perhaps as early as bears (Rogers 1976). Manville (1981) men- late December. Gestation lasts roughly about tioned depredations on grain fields, orchards, 210 days, but development occurs in the last beehives, and livestock. ten weeks. The young weigh only 225-330 g Two students of bear ecology in Wiscon- (7.9-11.6 oz) at birth, and are naked and blind. sin found that plants (grasses, sedges, berries) When young develop enough to leave the den, were the main foods eaten. An important food, in April, there is natural food available. The acorns, surprisingly, was not mentioned as young retain their milk teeth. These are re- important in either study. The common ani- placed when the bears are two years old. The mals eaten were ants. Norton (1982) studied young may stay with the mother even through 630 scats collected during May through Au- the winter of the second year. This care low- gust 1976 and 1977. Seasonally important ers infant mortality considerably. Upon emerg- foods were grasses, sweet cicely (Osmorhiza ing in spring the mother eventually drives the claytoni) and aspen (Populus tremuloides) in yearlings away, but sometimes they stay with spring; gooseberries (Ribes spp.), blueberries her up to 17 months until summer breeding. (Vaccinium spp.), and ants (Formicidae) in Many ages ago, Pliny the Elder, in His- summer; blackcherries (Prunus serotina), wild toria Naturalis, wrote this translated obser- sarsparilla (Aralia nudicaulus), and domestic vation, “Bears when first born are little shape- oats (Avena sativa) in late summer. Green less masses of white flesh a little larger than a vegetation comprised the bulk of the diet. There mouse, their claws alone being prominent.” was no mast (acorns and nuts) present in One of the truly remarkable things about any Norton’s study area (see Payne et al., 1998). kind of bears is the production of such tiny Bertagnoli (1986) analyzed 337 scats and young. To my knowledge, no other carnivore, found 45 kinds of foods (May-August). These perhaps no other mammal except marsupi- were collected in 1978 and 1979, and there als, shows such size limitation of the neonate. were differences in the foods eaten. Grasses Relatively tiny embryos are not the rule in and sedges were the chief foods. The plants large mammals, sexually promiscuous used most were in Rosaceae. Ants were the mammals, or predators. The bear is all most important animal food. Storlid (1995) three. The question arises, why does the bear

356 THE WILD MAMMALS OF WISCONSIN produce tiny young, being large, promiscu- Lyme disease), and a mite (Manville 1978; ous, and on occasion predaceous? The bear 1981; Addison et al. 1978; and Rogers sleeps extensively in winter, without a true 1975). The mite may cause mange in Wis- hibernation, to conserve the body fat, but it consin bears. Worms include Alaria, Baylisas- often emerges lean in the spring. Since the caris, Dirofilaria, hookworm larvae, and bear breeds in autumn, but has a delay in Physaloptera. Diphyllobothrium latum is implantation, the dormant, unimplanted em- now rare (Rogers, 1975) in bears. The inci- bryos cannot make use of metabolized ener- dence of Trichinella spiralis is low in Wis- gy from their mother’s winter body fat. Neo- consin and Michigan (Zimmerman 1977). nates gain nutrients (i.e., high in titre) from There are doubtless some respiratory and vi- the mother’s milk for rapid growth and devel- ral diseases that affect bears. More study is opment. It is good strategy for species with needed to identify them. dormant embryos to either conceive in spring Manville (1976) made a thorough study or bring forth tiny young as do the bears (Ram- of black bear parasites from bears in northern say and Dunbrack, 1986). Black bears live Wisconsin. Ectoparasites were collected from long lives, up to 25 years, but usually 10-15 nine bears captured in 1974, and 104 bears (Crandall, 1964). Bears may live and proba- (including 18 recaptures) taken in 1975. Vis- bly breed for as many as 13 years in the wild cera from 28 bears killed by hunters during according to Baker (1983). In Wisconsin some the 1974 and 1975 bear hunting seasons were have been aged well over 15 years. One age examined for internal parasites. Thirty-five of is mentioned as approximately 30 (Kohn and 44 bears from the vicinity of Clam Lake car- Rolley, 2000). The mean age is about four ried the dog tick (Dermacentor variablilis), and years for harvested bears. one the black-legged tick Ixodes scapularis— The number of young in black bears of a new record for bears. Forty-five of 60 bears Wisconsin is usually three (range 1-3). Matson from Iron County in 1975 carried dog ticks. (1952) reported a litter of five. The altricial Four Iron County bears taken in 1975, had young are only six to eight inches (150-200 mallophagan Trichodectes pinguis. Mange mm in length, weighing half a pound or per- was observed on five bears; two Demodex haps a pound, usually about 250 g each). The mites were found in a scab. A flea was found eyes and ears are closed, and there is no fur. on one bear from Iron County (1974). Fur appears on the back in a week, and at 25 Twenty-five intestinal tracts (89.3 per- days the eyes open. At 46 days, the ears open cent) contained ascarid worms (Baylisascaris). and the weight is over five pounds (2.27 kg). Fifty-nine of 92 fecal samples had the eggs. By then the upper incisors have erupted. Hookworm larvae were found in a bear caught Mortality. Aside from people hunting and in 1975. Adult filaria worms (Dirofilaria ursi) killing bears that are pests, the black bear has were found in two. Seventeen (19.5 percent) few enemies. Starvation and disease take the blood smears contained microfilariae. The very young (and probably the very old), where- worm Trichinella was found in only 6 of 163 as hunting removes bears one year old and (3.7 percent) bears (1970-1973). older. Mortality may be high in cubs and year- Dental caries are not uncommon in lings if autumn foods are scarce. Mast fail- bears, probably due to their eating sweet foods ures may effect a heavy mortality. The timber (E. R. Hall, personal communication). wolf may on occasion kill a small bear (Rog- Home Range and Density. Home range ers and Mech 1981). Cannibalism occasion- studies were made on the Upper and Lower ally occurs (Trauba, 1996). peninsulas of Michigan (Erickson 1964; Man- Parasites include flea, louse, several ticks ville 1980, 1981). Home ranges of Upper (including the tick, Ixodes, that transmits Peninsula black bears averaged 12,955 acres

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 357 (5,234 ha) for males and 6,477 acres (2,617 dens later than the females and young do, for ha) for females. In Lower Michigan, Manville winter sleep. On the Upper Peninsula the found more extensive wandering, 32,400 earliest date for denning was October 13. In acres (12,928 ha) and 16,550 acres (6,668 southern Michigan and in Wisconsin bears ha) for males and females, respectively. Abun- doubtless enter torpor later and sleep less, dance of bears is estimated based on harvest with dormancy perhaps irregular (Manville figures (tables Car-10-11). In 1995-1996, Kohn et al. (1998) found that young males made up the most of the harvest, through the Table Car-10. Age classes of bears harvested in Wiscon- first four years of life. Yearlings are most of- sin, 1986-96. ten killed. After the males are thinned out, Percent in age class more females are taken from the 4th to 10th Year 1-2 yr 3-9 yr 10+ yr No. aged Mean age year, or occasionally later for very old bears, 1986 Males 59.5 37.2 3.3 210 3.6 falling to less than 10 percent in either sex in Females 43.8 41.3 9.9 121 4.2 any of those years. The estimated densities 1987 Males 52.6 43.2 4.2 401 4.1 varied from approximately 1 per mile to low- Females 41.5 52 6.5 200 4.6 er values (1 per 6.3 square miles) according 1988 Males 60.4 35 4.6 439 3.7 Females 40.9 51.9 7.2 345 4.7 to Kohn et al. (1982, 1998). Weber (1994) 1989 Males 53.9 39 7.1 397 4.2 found 1/6.1 square km in northeastern Wis- Females 42.5 47.9 9.6 261 5 consin, in predetermined “good habitat.” 1990 Males 67 30.4 2.6 454 3.4 Movements by three bears, all males, in Females 46.8 48.1 5.1 331 4.6 Lower Michigan, were 65, 95, and 35 miles 1991 Males 56.9 37.3 5.8 448 4 Females 38.9 54.9 6.2 306 4.7 (104, 152, and 56 km, respectively) on un- 1992 Males 63.9 32.1 4 474 3.5 usual autumn excursions. These bears then Females 48.4 45 6.6 380 4.3 returned to their home areas by November of 1993 Males 50.9 41.7 7.4 405 4.3 the same year. Harger (1970) reported hom- Females 37.8 57.3 4.9 286 4.6 ing in a bear traversing a distance of 142.5 1994 Males 62.6 31.4 6 441 3.9 Females 50.9 45 4.1 271 4.2 miles (228 km). Some bears stay where they 1995 Males 55.7 41.4 2.9 600 3.6 were transplanted. Females 37.7 52 10.5 435 5.3 Remarks. Nowack (1991) ably reviews 1996 Males 60 37.3 2.7 771 3.6 the physiology using the term “hibernation”, Females 46.8 45.6 7.6 536 4.7 as do many lay people and some naturalists, but black bears do not hibernate. They do undergo a drowsiness or torpor the Germans Table Car-11. Number (%) of black bears harvested in call Winterschlaf. Winter-sleep is a good term Wisconsin by sex and age class, 1995-96 (After Kohn et al., for this condition. The great biologist, G. 1998) . Cuvier, in Regne animal, 1846, used the Age Class 1995 1995 1996 1996 meaningful French word for drowsy-sleep as (years) Males Females Males Females “le sommeil” or somnolency. [This word in 1 184 (30.7) 84 (19.3) 251 (32.6) 103 (19.2) French means drowsiness or sleep, and does 2 150 (25.0) 79 (18.2) 211 (27.4) 148 (27.6) not mean true hibernation.] The body fat in 3 126 (21.0) 71 (16.3) 104 (13.5) 58 (10.8) early winter is about 40 per cent of the body 4 46 (7.7) 34 (7.8) 84 (10.9) 73 (13.6) weight. The bears subsist on their fat and sleep 5 35 (5.8) 41 (9.4) 34 (4.4) 33 (6.2) unconcerned when people are nearby, but 6 11 (1.8) 28 (6.4) 34 (4.4) 27 (5.0) 7 17 (2.8) 26 (6.0) 17 (2.2) 23 (4.3) they may arouse violently when disturbed. 8 10 (1.7) 15 (3.4) 14 (1.8) 16 (3.0) Females often move their cubs to a different 9 4 (0.7) 10 (2.3) 2 (0.3) 15 (2.8) winter den if disturbed. Adult males enter their 10+ 17 (2.8) 47 (10.8) 20 (2.6) 40 (7.5)

358 THE WILD MAMMALS OF WISCONSIN 1980). The heart rate drops 8 to 10 beats per minute or even lower, but the body temperature drops only about 11 degrees F (3-4 degrees C). Summer temperature is about 100 degrees F (37 degrees C), and in winter it is about 90 degrees F (34 degrees C). In spring, an arousing bear defecates a hard and dry fecal “plug”. Bears may arouse thin or fat depending on their metabolism, the amount of autumn food, severity of winter, and so forth. Thin bears are said to be very hungry, but some emerging bears do not eat for several days. Additional natural history. Pelton (1982) and Nowak (1991) reviewed the natural history for the black bear. Geographic variation. There is but one race in Wisconsin and Upper Michigan. Specimens examined. Total, 10. Ashland Co.: Clam Lake 1. No specific locality 1. Door Co.: Beach, N end Washington Island 1. Lang- lade Co.: T34N, R9E, Sect. 19, Parish Twsp 1. Lincoln Co.: Hwy 29, N of Antigo 1. Mari- nette Co.: No specific locality 1. Portage Co.: No specific locality, probably Jordan Swamp 1. Price Co.: No specfic locality 1. Shawano Co.: Between Tigerton and Wittenberg 1. Wis- consin No specific locality 1.

Raccoon family. By T. Swearingen. Kansas University.

Family PROCYONIDAE Bonaparte suspended on a gold chain, and used as a The Procyonidae are found in Asia, and North toothpick. The raccoon in Wisconsin in re- and South America. Most members have an- cent decades has increased from rare to ex- nulated tails. The feet are plantigrade, al- tremely abundant also, in spite of several dis- though dextrous. The procyonids are found eases (see below) and popularity of its fur. This at home in trees or ranging about on the increase in numbers is true in other states. ground. The cheek teeth, reflecting an omnivorous diet, are bunodont, especially those behind the carnassials. Although the penta- Genus Procyon Storr dactyl feet resemble those of bears, and the cheek teeth are similarly bunodont, there is Procyon lotor (Linnaeus) no alisphenoid canal and the baculum (and Raccoon penis) is relatively large and unique. In Procy- on lotor the baculum is bifid distally, with “The upsurge in raccoon numbers, which has rounded terminations, and the end is hooked. occurred in many areas during recent years... is The base is robust. Years ago, as a rather surely a major irruption”. — Glen C. Sanderson, crass curiosity it was ground sharp on the end, 1951. [also in furs taken.]

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 359 1758. [Ursus] lotor Linnaeus. Syst. naturae, ed. 10. open the vagina, and hook into it during coi- 1:48. Type locality Pennsylvania (fixed by Th- tus (Long and Frank 1968). Rinker (1944) omas, Proc. Zool. Soc. London, p.140, 1911). reported an os clitoridis. There are six mammae, two pectoral, two abdominal and two The name Procyon means early or first inguinal. The chromosomes are usually 2N = dog, although Procyon is not an early dog at 38 (Lotze and Anderson 1979). all. Some say it means “before dogs.” The word The raccoon braincase is quite rounded. lotor means the washer, based on the sup- Occasionally there is a low sagittal crest. One posed and overstated habit of captive raccoons adult specimen (UW-SP No. 2761) has a well- that may wash their food in water before eat- developed sagittal crest. The auditory bullae ing it. According to Jackson (1961) Captain are inflated. The infraorbital canal is often John Smith described the raccoon as a kind of divided as two foramina on each side, at least badger called “aroughcun”, which came from in Wisconsin and several other states I the Algonquian Indian name arankun. checked. The upper third incisor has an ac- There is a single geographic race in Wis- cessory lobe. The molars are not sectorial, consin and it extends across Upper Michigan and may be called bunodont. The cheek teeth as well. Whether it ranges throughout the are subquadrate, about as broad as long. The Lower Peninsula requires a quantitative ap- upper carnassial is somewhat sectorial. The praisal, but previous simplistic assignments by first upper premolar has a single root. There Jackson and Burt, respectively, resulted in one are ten teeth in any row, provided there is no of the previously discussed Jackson-Burt divi- dental anomaly. The hard palate on the skull sions, P. l. hirtus for Wisconsin and P. l. lo- extends far back, the distance from pterygoid tor for Michigan. The specimens I have seen to the back molar about equal to the length both in Wisconsin and Upper Michigan are of the maxillary tooth row. Anatomical stud- referable to P. l. hirtus. ies on raccoons are summarized by Lotze and Anderson (1979). The raccoon shows a prominent brown- Procyon lotor hirtus Nelson and Goldman black facial mask set against adjacent whitish hair. The black hides the carnivore’s bright 1930. Procyon lotor hirtus Nelson and Goldman. eyes, and the black is continuous with a medi- J. Mammalogy, 11: 455. Type from Elk River, al black band from the black nosepad to the Sherburne Co., Minnesota. level of the ears. They are edged with white. The tail is fairly long and annulated with five to Description. This medium-sized carni- seven blackish bands alternating with pale gray- vore, aside from its distinctive color pattern ish rings. The fur is grizzled, heavy on the black (mask, ringed tail) is recognizable because of and iron gray, but occasionally brownish, red- the broad head, narrow muzzle, pointed ears, dish brown, or yellowish. The underparts have and bushy tail. There are five slender toes on brownish underfur overlain with gray or brown each foot, each with a long claw, and the feet guard hairs. There are numerous color vari- are nearly plantigrade. The soles are naked. ants known to breeders, and golden red, mel- The raccoon often sits on its hind feet and anistic, yellowish, and albino are the most com- manipulates or searches for food with its mon. Long and Hogan (1988) found that albi- hands. Often it does this in water. The gait of nism in raccoons results from two different a running raccoon is peculiar, with the legs recessive alleles, and that albino parents hav- extended and the back arched high. The bac- ing different recessive alleles may by epista- ulum is long, hooked, and distally bifurcate, sis produce normal colored offspring. Both terminating as two condyle-like structures that these albino phenotypes had pink eyes, but

360 THE WILD MAMMALS OF WISCONSIN the golden red mutant had black eyes. The 38.8, 36.9, 38.1/ 41.5, 35.3; zygomatic annual molt lasts through most of the summer breadth 77.5, 75.2, —, 74.8/ —, —; inter- (Goldman 1950, Steuwer 1942), in Michigan orbital breadth 24.3, 25.3, 23.6, 23.6/ 24.9, from April to August. Wisconsin specimens 22; maxillary toothrow 41.9, 43.6, 41.9, showed molt September through late Decem- 43.5/42.3, 44.4. ber and prime fur into April. Dental formula. I 3/3, C 1/1, P 4/4, M Adult raccoons average 14-24 pounds 2/2 = 40. Teeth may be missing or supernu- (5.6-9.6 kg) but may reach 40 pounds (18 merary (Goldman 1950). kg) (Bluett and Craven, 1985). Raccoons Geographic Distribution. The raccoon is reach a meter (39 in) in total length. Cranial found in every Wisconsin county and often in measurements of 4 male adults and 2 female urban and cultivated areas. It is distributed in adults are: Condylobasal length 112, 118, diverse habitats too, from hills to low swamps, 114, 113/ 114, 115; length of nasals —, and is found on Washington and Rock islands in Lake Michigan. In Wisconsin, raccoons are abundant along the St. Croix River in the northwest, in southern Wisconsin, and northeastward to Rock Island. They are least abundant in northern counties, unknown in the Apostle Islands, but widespread in Upper Michigan. The raccoon, which prefers hardwoods, was neither wide-spread nor common on the Upper Peninsula until late in the 1930’s (Baker 1983). Records of specimens of raccoons examined and other reports by Jackson and myself practically blanket the state, with the exception only of Shawano, Iron, and Florence counties. A recent study by Amundson and Marc- quencski (1985) used carcasses supplied by the Wisconsin Trapper’s Association, with recent records statewide except along the southern counties. Iron County was included. Status. The raccoon is doubtless the most abundant wild carnivore in Wisconsin (Table Car-1), approached in numbers only by the striped skunk Mephitis. Attention to road-kills in Wisconsin in any summer will confirm this conclusion. For example, in July 1998, I observed 1 or 2 dead young raccoons in every county from Redwing, Minnesota, to Apple- ton, Wisconsin, mostly along Highway 10. Continuing eastward, raccoons were also seen dead in Brown and Door counties. In 1999, the same extensive road-kill mortality was observed, in late July and early August. In 1999-2000 the harvest was estimated as Skulls of Procyon lotor. Dorsal and ventral views No. 2186 young, Dorsal 2761 Old with sagittal crest and fused 130,141 (Dhuey, 2000), mostly from Clark, nasals. Jefferson and Sauk counties.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 361 The raccoon is managed throughout Wisconsin by hunting and trapping regulations. The old Wisconsin Conservation De- partment once (in 1936-1950) introduced raccoons into numerous Wisconsin counties; thousands were propagated and released. When this program was terminated the raccoon surprisingly continued to increase. The peak harvest was about 250,000 in 1978- 1979, and the peak price surprisingly lagged two years (1979-1980). The raccoon is aesthetically popular with people, and is seldom killed by them even when destructive in the

Maps showing geographic distribution of Procyon lotor in Wisconsin and North America.

362 THE WILD MAMMALS OF WISCONSIN garden. Its supposed habit of washing food, on McDill Pond). They may sleep in woodpiles, well known antics as an adopted pet (but see hay stacks, and ground dens made by other beyond), appealing “hands”, bright eyes, and mammals (Lotze and Anderson 1979). Some- soft fur are endearing, even when it overturns times numerous raccoons hole up together for the garbage cans, digs up the lawn for acorns winter (Mech et al.,1966). or grubs, steals the dog’s food, or eats the Foods. The raccoon feeds extensively on farmer’s sweet corn and other produce. plant parts, especially fruits, and on a diverse When corn sweetens (the so-called “milk” diet of animals as well. Raccoons have fed on stage) raccoons may ravage an entire field. crippled waterfowl (Yeager and Elder 1945), For large problems the U. S. Department of turtles and their eggs (Erickson and Scudder Agriculture’s Wildlife Services can be notified. 1947), herring gull eggs and young in Massa- A large live trap can trap out raccoons, which chusetts (Kadlec 1971), some other birds, and may be transported to a remote place (sug- occasional reptiles, frogs, rarely toads and gested as at least 50 miles away). salamanders (Kaufmann, 1982). In Illinois Not so happily for the raccoon, its fur is (Yeager and Rennels, 1943) foods include fox luxuriant and valuable. The meat is dark and squirrel, muskrat, cottontail, mice, shrews, fatty but edible, and “running” raccoons with ducks, cormorant, chicken, fishes, snails, hor- dogs (“‘coon” hunting) is a popular sport at nets, water bugs and beetles, crayfishes, per- night. To hear the cry of hounds on the track simmon, pecan, grapes, pokeweed, and mis- of a raccoon or opossum, and the baying of cellaneous items, opportunistically eaten in the the dogs when the raccoon is treed in the autumn (October to November). Jackson moonlight of a frosty autumn evening is an (1961) listed for Wisconsin diet a preponder- experience not soon forgotten. In Wisconsin ance of plant foods such as black cherry, there are a few raccoon breeders, who espe- chokecherry, pin cherry, currant, wild grape, cially hope for albino raccoon fur (Long and hawthorn, wild plum, apple, tomato, black- Hogan 1988). berry, raspberry, strawberry, blueberry, and In 1993-1994, about 160,000 raccoons other fruits and acorns, hickory nuts, hazel- were harvested in Wisconsin. Total pelt value nuts and beechnuts, field and sweet corn, oats, ($1,321,000) outranked all furbearers (Dhu- and other shoots and buds of plants. Animal ey, 1995). Between one and two million are foods are as much as 70 per cent of the food harvested each year in the United States. volume, particularly in diets including cray- Habitat. The adaptable raccoon lives in fish, snails, and insects, and contain about 30- varied habitats, preferring hardwoods with 40 percent plants. These animals include cray- numerous hollow trees for security and den- fish, snails, clams, earthworms, grasshoppers, ning. It seldom ranges far from standing wa- crickets, beetles, wasps, bees and moths, fish- ter. Hoffmeister (1989) suggests 1,200 feet es (especially bullheads), frogs, turtles, eggs (nearly 400 m) as a limit. Its foods show how and young of birds on occasion, mice, shrews, much it depends upon lakes and streams. muskrats, and squirrels (rarely). Dorney (1954) Raccoons may be found in treeless prairies found crayfish and young muskrats were eat- or swamps having no hollow trees. They are en in a Wisconsin marsh, along with corn, often found in urban areas. berries, waterfowl, and a few muskrats dis- The raccoon reportedly does not build a abled by humans. Some animal foods of the nest, but it may line a tree cavity with leaves. It raccoon are carrion. opportunistically dwells with its young in rocky Reproduction. Raccoons of both sexes crevices (observed in Wood County, Sand Hills may breed in their first or delay to the second Demonstration Area), attics of abandoned year. A female may have a second litter on farmhouses, and hollow snags (Stevens Point occasion (Bissonette and Csech 1938, Johnson

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 363 1970, Lehman 1968), but this is doubted by wer 1943, Johnson 1970, Stains 1956, Sanderson (1987), is likely rare, and probably Whitney and Underwood 1952). About unsuccessful. Late litters are not likely to sur- 200,000 per year were harvested in Wiscon- vive in areas as far north as Minnesota (Mech sin (Table Car-1) by hunters and trappers et al., 1968) and including much of Wiscon- (Bluett and Craven, 1985). In 1989, over sin. Some yearling females may not conceive 140,000 were harvested, and in 1991-1992 because of extremely cold weather, until their over 74,000 were trapped. In Wisconsin, second year (Stains 1956). In Root’s (1981) automobiles also cause much mortality (Root study in 13 southwestern counties in Wiscon- 1981). Mortality in the raccoon is greatest in sin, only 32 percent of the yearling females the juvenile age class. conceived. He found some juveniles (10 per The value of the raccoon fur leads to high cent) resulted from late litters conceived from human-caused mortality, once estimated at an mid-May through mid-July. Root (1981) found incredible 98 percent (43 percent hunting, 55 no juvenile males capable of breeding the first percent trapping) in southwest Wisconsin year. Root thought his litter size of 3.71 in (Root 1981). Tom Howard (personal corre- southwest Wisconsin resembled that of Mani- spondence) suggests this percentage must be toba (4.1, Cowan 1973) and North Dakota considered to include natural as well as hu- (4.5, Fritzell 1978), but I suggest this may in- man caused mortality, because no population deed be a significant difference, as the means can sustain 98% mortality. were based on adequate samples. The bobcat, fisher (Seton, 1953), red fox, Breeding takes place in winter (February- coyote and several owl species are known to March), and gestation is about 63 days (Sand- prey on raccoons (Stains 1956, Whitney and erson and Nalbandov 1973). Birth occurs in Underwood 1952). Starvation is suspected as late March, April, and May, and even later (Bak- a cause of death. Food shortages, diseases, er 1983). The newborn raccoon has blackish and parasitism all may lead to mortality (Lotze skin covered with fine grayish fur, an indistinct and Anderson, 1979). black facial mask, and tail rings alternating Raccoons carry diseases and have been whitish and black. The eyes and ears are closed. studied a great deal as indicator species for These open in about three weeks. Lower inci- monitoring pollutants. Knuth (1979) studied sors and canines also erupt by three weeks. At polychlorinated biphenyls (PCB’s) in raccoons nine weeks the young raccoon eats solid food. living near the Wisconsin River west of Plover, By 12 to 16 weeks the young is weaned and in Wisconsin. Often found in the semi-aquatic weighs about 3 to 4 pounds (= 1.4-1.8 kg). In mink, the presence of this dangerous substance four or five months permanent teeth replace in raccoons is, at the least, surprising. the deciduous teeth (Baker 1983, Hamilton Raccoons carry at least 13 pathogens 1936, Montgomery 1964, Stains 1956, and causing disease in humans (Bigler et al. 1973). Stuewer 1943). The mother often moves her Except for canine distemper (Robinson et al. litter from one den to another. 1957) and parvovirus the diseases do not Age of raccoons can be determined by seem of major importance in raccoon mor- cranial sutural closure, ossification of epiphy- tality. Parvovirus recently and sporadically ses, cementum layers of teeth, the size and killed many raccoons, as well as dogs, in Wis- extensibility of the baculum, and eye lens consin. Rabies and tularemia may be suspect- weight. Raccoons have lived 12 years in the ed in raccoons, but these deadly diseases are wild and 22 years in captivity (Haugen 1954, not prevalent in Wisconsin raccoons. I knew Crandall, 1964, Jones 1979). a fur breeder in Illinois raising raccoons ex- Mortality. The greatest enemy is human- clusively, who caught Lyme disease from ticks kind by means of hunting and trapping (Steu- associated with his captive raccoons. Some

364 THE WILD MAMMALS OF WISCONSIN raccoon diseases may be transmitted by the 13 acres (5.2 ha) and as large as 930 acres raccoon’s urine or feces. (372 ha) with an average of 268 acres (107 Distemper, pneumonia, and parasite in- ha). Fisher (1977) found raccoons to move festations periodically decimate American rac- distances of up to 1,720 feet. In Minnesota coon populations (see Lotze and Anderson one super-wanderer travelled 165 miles (264 1979). The protozoan Giardia and the round- km) (Priewert 1961). worm Baylisascaris may be contracted by In good habitat for raccoons, densities humans from raccoons. In southern Wiscon- varied from 1 raccoon per 10 acres (4 ha) to sin 50-77 percent of raccoons may harbor 1 per 16 acres (Dorney 1954, Yeager and this roundworm. In northern counties the in- Rennels 1943). Steuwer (1943b) calculated cidence is less (Amundson and Marcquenski, densities of 1 raccoon per 16 acres (6.4 ha), 1985). Ingestion of aged (about 30 days) eggs but in Clinton County Michigan, Linduska of this roundworm from the environment (1950) found only 1 raccoon per 43-55 acres. causes the infection. Baylisascaris can infect Bluett and Craven (1985) report densities as humans via the eggs passed in raccoon scats. 1 to 30 or 40 acres (12 to 16 ha) in Wiscon- The parasite occasionally causes death in small sin. In urban areas the density may be as high children and a rabies-like sickness in wood- as one per 12-20 acres (4.8-8 ha). chucks, tree squirrels, and rabbits. In Wiscon- Remarks. Raccoons are often taken from sin 51 percent of 213 raccoons from 43 coun- nature for pets. They may grow up to become ties contained this worm. Juveniles show high- aggressive and destructive. They cannot be re- er incidence than adults (Amundson and Marc- turned to the vicissitudes of nature and survive. quenski, 1985). An adult specimen from Dane County Jackson (1961) lists parasites including bit- (No. 109) has two sets of frontal bones. Some ing and sucking lice, ticks, and fleas. Internal skulls have uncommon, intricate (fractal) su- parasites include roundworms, Trichinella, tures (Long and Long, 1992), greatly enhanc- tapeworms, and one trematode (Clinostomum). ing their strength.. Home Range and Density. Males have The species is famous for occasionally larger home ranges than do females (Lotze washing its food in water. Actually, this has and Anderson 1979). Steuwer found in Mich- not been observed in nature. In studies on igan that adult males had home ranges as small albino raccoons I never heard that any cap- as 45 and as large as 2,000 acres (18 to 800 tives do it. ha, respectively), with an average of 500 acres A raccoon might share a den with the (200 ha). Females had territories as small as striped skunk (Shirer and Fitch 1970). They

Raccoon. By Walter Weber. Raccoon harvests (1,000) average pelt price. 1973-1983.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 365 may aggregate in a group for winter sleep- worth, Washburn, Waukesha, Waupaca, ing. Essentially nocturnal, raccoons are curi- Winnebago, Wood counties. ous and playful. They swim well, and can usu- Other Records (Balliett and Taft, 1978). ally best a dog in combat. The female can Monroe, Ozaukee, Trempealeau, and Sha- frequently be found with young by hearing wano counties. Personal observations of dead her twittering purrs. Raccoons do not truly on roads: Pierce, Buffalo, Pepin, Clark, Wood, hibernate, i.e., vary their body temperature Portage, Waupaca, Ozaukee, Brown, Door with cold ambient temperatures to any great counties. Most counties in 1985, Amundson extent, but they are often dormant or torpid and Marcquenski. for periods of several days in cold weather conditions. In Wisconsin, to my knowledge, the raccoon in winter has not been studied. Family MUSTELIDAE Swainson Additional Natural History. Lotze and Anderson (1979) reviewed natural history for This family includes the martens, weasels, the raccoon. mink, skunks, river otter, and badger. There Geographic variation. There is one race in are few placental families as diverse as this one, Wisconsin, northern Illinois, and Upper Michigan. and some workers suggest that differentiation Specimens examined. Total, 39. Adams, has been progressing for many millions of Dane, Door, Eau Claire, Iowa, Juneau, years. Perhaps the family should be divided Langlade, Lincoln, Manitowoc, Marquette, into two or more families. Recently Dragoo Oneida, Ozaukee, Portage, Rock (lact. Oc- and Honeycutt (1997) proposed that the tober 15), Rusk, Sheboygan, Vilas, Wal- skunks (Mephitinae) and the stink badgers (My- daus) should be recognized as the family Me- phitidae, in my opinion a premature opinion. For one example, no one has even described the penis and baculum of Mydaus. The crux of the problem is that the paleontological and anatomical evidence suggest badgers are a natural group, allied to one another fairly closely with a good fossil record to prove it (Long and Killingley, 1983; Long, 1881; and Petter, 1971). I suspect the resemblance of Mydaus and skunks is evolutionary convergence. Fur- thermore, skunks and otters are definitely related closely (Hunt, 1974; Wozenkraft, 1989; Wyss and Flynn, 1983) . On the other hand, those working with molecules and dendrograms find parsimonious arrangements suggesting skunks and Mydaus are much more closely related than the latter is to the other badgers. I submit some older evidence that was either overlooked or ignored by workers (Long, Figure showing the status of Bayliscaris procyonis in 1981; Petter, 1971), who in these and other Wisconsin. In Wisconsin, 51% of 213 raccoons from 43 works discussed subtle and derived characters counties contained this worm. In northern counties the tying the badgers together, with Meles and incidences is significantly lower than in the south, where the raccoons are abundant. Juveniles show higher incidence than Taxidea not so closely as supposed, and men- adults. After Marcquensky. tioning peculiar articulations of the teeth and

366 THE WILD MAMMALS OF WISCONSIN even the mammary gland arrangements sug- large and lined with pale hairs, marks of gesting common origin. The least division I yellow or white confined to chin, throat could accept at this time would recognize a and head ...... Martens...4 family (or subfamily) of otters, skunks, and bad- 4 Size large (up to 1.0 m in males), fur gers (including Mydaus with badgers). I point brown with legs, back and belly often out that the parsimonious dendrogram on mo- blackish, hairs grizzled and coarse ap- lecular evidence given by Dragoo and Honey- pearing, throat lacking orange or white cutt, conflicts with strong morphological evi- marks, ear lined with whitish hairs, tail dence, lumps stink badgers with skunks, plac- longer than 290 mm ...... Fisher es Meles and Taxidea close together, and that Martes pennanti the family Procyonidae stood closer to them 4’ Size medium (up to 0.6 m, approximately than to the badgers Mydaus. In this work I two feet total length), color yellowish or follow the conventional use of the name Mus- reddish brown, head often grayer or yel- telidae, noting only that there is a strong affin- lower, feet and tail darker, fur fine and ity of the mustelines to one another. soft, throat and chin with orange or buff marks, tail shorter than 290 mm ...... American Marten Key to Wisconsin Mustelidae Martes americana 4’’ Size medium, color brown with white Characters of Skull and Body markings on throat and head, fur fine 1 Dental formula 36 teeth, upper molar and soft, tail shorter than 260 mm ...... rounded posteriorly and sulcate on oc- ...... Stone marten clusal surface, with high labial margin, Martes foina. Non-native auditory bullae flattened, hind feet introduction SE Wisconsin webbed, tail stout and furred (not bushy) 3’ DF 34 teeth, with three upper premo- ...... River otter lars, ear pinna medium or small and lined Lontra canadensis sparsely with dark hairs, markings not 1’ Dental formula not 36, upper molar not confined to chin, throat and head (ex- rounded much inward and posteriorly, cept in the mink, which has tiny, obscure not sulcate with high labial margin (ex- ear pinnae) ...... 5 cept in skunks), auditory bullae not flat- 5 Upper molar sulcate as in otter, much tened (except in skunks), hind feet never larger than upper carnassial, auditory webbed, tail bushy ...... 2 bullae flattened (as in otter), color glossy 2 Upper molar and carnassial both trian- black with bold white stripes ...... gular, skull wedge shaped, broader pos- ...... Skunks 6 teriorly, forepaws and foreclaws enor- 6 Size medium (approximately 0.6 m, mous, black “badges” on cheeks and a about two feet total length), white stripes short stripe on head and nape of neck . variable, dorsolateral and longitudinal and ...... North American Badger often joining near the neck ...... Taxidea taxus ...... Striped skunk 2’ Upper molar “dumbbell” shaped* or Mephitis mephitis deeply dished (in skunks), skull not 6’ Size small (approximately 45 to 55 cm, wedge-shaped, forepaws ordinary and 18-21 inches total length), white stripes not specialized for digging, black “badg- several, often broken and extending on es” and head stripe lacking ...... 3 the neck, shoulders, back and sides ..... 3 Dental formula 38 teeth, with four up- ...... Spotted skunk per premolars on either side, ear pinna Spilogale putorius

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 367 5’ Upper molar “dumbbell” shaped *, audi- * A “dumbbell” is an antique weight expand- tory bullae moderately inflated, color ed at either end used for exercise of the arms. brown above ...... 7 ** Extinct in Wisconsin, may be found as 7 Size large (total length about .6 m, or subfossils. two feet), prominent sagittal crest, color *** The reader is advised to consult the de- all brown except for yellowish tan stripes scriptions herein of these weasels, which often laterally, some obscure yellowish mark- present great difficulties in their identification. ings on chest ...... Wolverine Gulo gulo** 7’ Size medium or small (total length up to Genus Martes Pinel .6 m), sagittal crest not prominent, bi- Martens, Fisher and Sables colored brownish aboveand yellowish or whitish below, except in winter when the Martes is the most primitive genus of the Mus- pelage is mostly white (except in the mink telidae, except possibly the Oriental ferret bad- which is brown below, and never white gers, having 38 teeth (more than the advanced in winter, see couplet immediately fol- mustelids, which have 34 or 32), sharp car- lowing) ...... 8 nassials, anal scent glands, slender body and 8 Total length 0.45 m (18 inches) to 0.6 fairly short legs, and an upper molar which m (two feet), all brown except blackish seems generally ancestral to the dumbbell in tail and white markings on chin and shape of Mustelines and of the broad occlusal throat, ear pinnae small ...... Mink molar forms of badgers, otters, and skunks. Neovison vison This upper molar is somewhat constricted an- 8’ Total length up to .43 m (17 inches), teroposteriorly, but there is a relatively large brownish above and tan or whitish be- heel on the lingual expansion. There are large low, except in winter white pelage, ear auditory bullae, and the profile of the skull is pinnae medium ...... Weasels 9 flattened, with stout rostrum almost in line with 9 Size small (up to 206 mm total length, the cranium. The skull, especially in old males, 7.5-8 inches), tail short (approx. 38 mm), often has a prominent sagittal crest. The males hardly black-tipped (showing a few black are larger than the females, but in anterior fea- hairs), skull approx. 30 mm length (less tures of the cranium and in the carnassial teeth than 1.5 inches) ...... Least weasel the females are not much smaller, indicating Mustela nivalis the prey is not much different and prey selec- 9’ Size longer than 210 mm total length, tion did not lead necessarily to sexual dimor- tail medium or long, tail distinctly tipped phism (see Holmes and Powell, 1994). There with glossy black, skull longer than 34 are only four mammae, all inguinal, a very prim- mm ...... 10 itive pattern in Mustelidae. 10 Tail over one third total length, usually Martens tend to be slender, adept climb- 40 percent, with long black tip, rostrum ers in trees or quick scamperers on the ground of skull elongate, postorbital processes enabling them to catch birds and rodents. One in adults prominent and pointed ...... functional adaptation is the ability to turn the ...... Long-tailed weasel hind limbs and toes about, as in bats, to resist Mustela frenata*** gravity and allow them to easily run down tree 10’ Tail less than one third total length, with trunks head first. shorter but distinct black tip, rostrum See Buskirk, Harestad, Raphael and short, postorbital processes in adults Powell (1993) for an account of the martens. hardly protruberant or tiny ...... Ermine Powell (1981, 1982) wrote two thorough Mustela erminea*** works on the fisher. The martens tend to re-

368 THE WILD MAMMALS OF WISCONSIN produce with a delay in the implantation of Thomas Pennant. The name fisher has no embryos, as follows, respectively, for Martes known source, but probably comes from the americana, M. foina, and M. pennanti: American Dutch name fishet (and related Duration Birth forms of the word) which means polecat in Aug.-Feb. Mar.-Apr. Dutch, and is the name of the polecat pelt in Aug.-Jan. Mar.-Apr. French. The original meaning of the word fish- Apr.-Feb. Mar.-Apr. et was “nasty animal”. (After Mead, 1994) Description. The fisher is a medium-sized carnivore with slender build, but stocky in comparison to Mustela. The fisher is the larg- Martes pennanti (Erxleben) est member of the genus Martes (twice as Fisher large as the marten, and about four times as heavy. It resembles the American marten M. 1777. Mustela pennanti Erxleben. Systema regni americana, but is larger, darker brown (silver animalis... p. 470, type from eastern Canada tipped or grizzled on the shoulders), lacks an (=Quebec). orange or buffy ochraceous splotching on the 1912. Martes pennanti pennanti: Miller. Bull. U.S. chin or throat, has instead irregular white Nat. Mus., 79: 94. blotches, has relatively short ears, and on the 1777. Mustela canadensis Schreber. Die Saue- skull the upper carnassial has an exposed la- gthiere... 3: 492. Type is Pekan of Buffon. bial root. The labial length is greater than 9.5 1784. Mustela melanorhyncha Boddaert. Elenchus mm, and the length of the carnassial molar animalium, p.88, type based on Pennant’s fisher. exceeds 11 mm. The tail is close to 300 mm 1800. Viverra piscator Shaw. General zoology... 1: (12 in) in length. The related skunks are bold 414, type based on Pennant’s fisher. black and white in pelage color, with flattened 1802. Mustela nigra Turton. A general system of na- auditory bullae, less constricted upper molar ture..., p. 60. Type based on Pennant’s fisher. (anteroposteriorly), and sharper angle of the 1829. Mustela canadensis alba Richardson Fauna Bo- rostrum and cranium outline (face). They are reali-Americana, P. 54, type from Hudson Bay. usually smaller than fishers. Mink are usually 1829. Mustela godmani Fischer. Synopsis mammalium, smaller and have shorter fur and much small- p. 217, type based on Godman’s M. pennanti. er ears. The fisher has shaggy fur, and about 1952. Mustela permanti permanti [printer’s erro- the head and shoulders it has tricolored hairs, neous spelling of pennanti] Barger. Wiscon- which give a hoary tan or silvery whitish wash sin mammals. Wisconsin Cons. Dept. See Jack- to the pelage. The feet are pentadactyl and son, 1961: 333.

Martes pennanti pennanti (Erxleben) See synonymy above

“its intestine contained hundreds of porcupine quills arranged like... packages of needles... passing along its interior as smoothly and surely as if within a tube of glass or metal.” – C. Hart Merriam, quoting N. A. Comeau, 1886.

The name Martes means marten in Lat- in, and pennanti honors an early naturalist Photo of mounted fisher in UW-SP Mammal Collection.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 369 semi-plantigrade, with semi-retractile claws lighter brown, almost chestnut or golden and furry soles of the feet. Leach (1977a, brown. Some are grayish, and some mutants 1977b) and Leach and de Kleer (1978) care- are white, fawn, or mottled white (Jackson, fully described the skeleton and forelimb mus- 1961). There are white blotches ventrally near cles of both fisher and American marten (Mar- the groin and the axillary areas, as well as on tes americana). The skull has a flattened ap- the chin and throat. There is an autumn molt pearance, with the rostrum well developed, to prime fur. Guard hairs attain lengths of and the snout, in external appearance, is rath- about 70 mm on the back. er long and fox-like. The fisher is significantly larger than the The baculum in young males is hooked American marten, without observed overlap and perforated with a foramen distally. In early in specimens of known age. This disparity is development the foramen was open distally. seen in external and cranial measurements. In adults the baculum develops a massive base Males are 15 to 18 percent larger in linear with prominent ridge (Wright and Coulter, dimensions than females, and much heavier. 1967); its length usually exceeds 10 cm. Males exceed 90 to 100 cm (40 in) in total Amorphous bone is deposited distally on old- length, and females do not reach that length er bacula. (Jackson, 1961; Powell, 1981). Males weigh The sagittal crest of the cranium projects up to 5.5 kg, and females as much as 2.5 kg. posteriorly in old males. To a lesser extent Cranial measurements in mm for two speci- the crest forms in females. The zygomata are mens (male and female, respectively) in the slender and weak (Powell, 1981) as are the UW-SP Museum are greatest length 121.5, attached masseter muscles. The karyotype is 104; condylobasal length 117.5, 100.9; max- 2N= 38 chromosomes (Benirschke and illary tooth-row 42.3, 37.8; zygomatic breadth Young, 1966). There are six inguinal (lower 65, 55.4; and interorbital breadth 25.6, 22.6. abdominal) mammae. Dental formula. I 3/3, C 1/1, P 4/4, M Fishers are dark brown, becoming black- 1/2 = 38. In comparing with mink or skunk, ish on feet and posterior (rump and tail). The the count of four premolars in any row iden- grizzled brown, even tan, on the shoulders tifies Martes. The long axis of the upper molar and head often appears grayish-tan. The eyes is almost perpendicular to that of the upper are dark brown or black. Some fishers are carnassial, and the inner cusp on the upper carnassial is distinctly set off from the main portion of the shearing tooth. The upper cheek teeth are larger than in the marten, the carnassial exceeding 9.5 mm. Distribution. Formerly statewide in forested areas, the fisher was restricted to forested areas in a few northern counties by the 1920’s (Jackson, 1961; Schorger, 1942, Scott, 1939). It probably has been extirpated since then, but was reintroduced (with plantings from New York and Ontario (Petersen et al., 1977)). Today, fishers are thriving and expanding their geographic range away from the release sites. There are many records in the Nicolet and Chequamegon National For- ests, in surrounding counties as well, and in Skull of Martes pennanti. upper Michigan. One record is in Marathon

370 THE WILD MAMMALS OF WISCONSIN Maps showing geographic distribution of Martes pennanti in Wisconsin re-established in Wisconsin and North America.

County, and I have a specimen, which seems 60) were brought from Minnesota and re- doubtful and was probably moved by human leased in the Chequamegon National Forest hands, from near Appleton. There are claims (1966-1967), in Ashland and Bayfield coun- of a specimen from Fort McCoy. In winter, ties (Pils and Martin, 1985). A limited trap- 1999, a specimen was obtained from a car- ping season was opened in 1985. Two areas cass found on McDill Pond, in Portage Coun- closed to trappers were established where only ty, where I have lived without fishers many wet trap sets were allowed. These were years. Obviously the species is spreading into 120,000 acres in the Nicolet National Forest new counties. See map. and 220,000 acres in the Chequamegon Status. See Distribution above. In the National Forest. By 1974, the fisher was early 1900’s fishers were regularly trapped known from 11 counties, and now from many in northern counties (Jackson, 1961), and more (See map). were taken and observed until the 1920’s and In Upper Michigan, a similar story is told, 1930’s (Jackson, 1961, Schorger, 1942; and the successes in both states became one Scott, 1939). This furbearer was apparently confluent population of fishers. According to extirpated, with no valid records (some otters Baker (1983) the U.S. Forest Service and the were trapped and reported as fishers) after Michigan Department of Natural Resources the 1930’s. Reintroductions are the likely obtained 61 fishers from Minnesota, New source of all new populations, which under York, and Ontario for release in the Ottawa management and legal protection are doing National Forest in the early 1960’s. The fish- well today. The original reintroduction (1956), ers were released in three different regions, into a 40,000 acre management area in the and by 1976, fishers were found in all the Nicolet region, consisted of seven fishers northern counties of Upper Michigan, and as (1956-1957), followed by another seven far eastward as Marquette and Delta counties (1957-1958) from the Adirondacks. In the (Baker 1983). The fisher was eradicated by next decade (until 1966) 120 fishers were lumbering off the forests and heavy trapping brought to Wisconsin from Ontario and New for its valuable fur. The successfully reintro- York (Petersen et al., 1977). Some (at least duced fisher is a fur animal, legally trapped in

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 371 certain northern counties. The female, al- and forest edge. In Maine (Arthur et al., though smaller, has a softer, more valuable 1989a) the fisher spent much of the time rest- fur. In 1994, fisher pelts sold for $12.50- ing in conifers. Dense conifer forest is not 58.40 per pelt, average $27. In 1992-1994, preferred habitat. Besides food, there must harvests exceeded 1,500 per year (Kohn and be mature deciduous trees, with hollows and Ashbrenner, 1995). A high harvest (3,644 cavities, which are used for the natal dens. taken) during the 1997 season reduced the Although the fisher will live in ground bur- state poppulation about 25 percent, but in rows in winter, it raises its young in Wiscon- 1998 a smaller harvest (496 fishers taken) sin in hollow trees (Jon Gilbert, personal cor- seemed to increase the numbers again (Kohn respondence, Coulter, 1966; Hamilton and et al., 1999). In 1999, approximately 721 Cook, 1955; Gilbert, 1995). Beliefs that the were taken (Dhuey et al., 2000). fisher requires closed canopy are untrue, but The fisher is beneficial, a valued furbear- wetlands support few hollow trees. er preying primarily on mice and hares, but it Most homes used by fishers are tempo- kills a few birds. Some consider predation on rary dens (Powell, 1981), but the fisher may hares and porcupines beneficial to those pop- return to them more than once. Such dens ulations, removing the sick and weak. The are hollow trees, logs, and stumps, brush piles, increase in fisher might effect a decline in rocks, abandoned beaver lodges, and holes porcupines. in the ground or snow. Habitat. Fishers in northern Wisconsin Foods. Foods vary with the habitat and usually inhabit closed-canopy and continuous the abundance of prey. Snowshoe hares, forest, but upland forests had the highest use squirrels, red-backed voles, and meadow voles whereas lowland conifer and shrub habitats are favorite foods. In some places porcupines were avoided. Diversity of forest types (hard- and raccoons make up much of the diet. As woods, second growth) provide a prey diver- mentioned above, birds and carrion of deer sity leading to an abundance of fishers (de are important foods (de Vos, 1952; Hamil- Vos, 1951, 1952). Tree species on the up- ton and Cook, 1955, Powell, 1979). In North lands include birch, quaking aspen, balsam America insects make up to 45% of the diet fir, and white spruce. Basswood, hemlock, in some places, vegetation varies up to 65%, ironwood, and white oak were present. Low- and birds are important in half of the studies land hardwoods included American elm, black (Buskirk et al., 1993). Occasionally the fisher ash, and maple. Near streams alders and wil- caches the kill for later meals. Fishers eat, in low were common (Kohn et al., 1993). In addition to the aforementioned foods, star- addition to forestland, fishers must have avail- nosed mole, masked shrew, water shrew, able prey, especially small and medium-sized short-tailed shrew, eastern cottontail, wood- mammals and birds and large carcasses of deer chuck, eastern chipmunk, flying squirrels, or other mammals. See Foods below. beaver, deer mouse, tree squirrels, white-foot- In contrast to the American marten (Mar- ed mouse, muskrat, meadow jumping mouse, tes americana) the fisher may inhabit forest marten, mink, striped skunk, ducks, ruffed edge and cut-over woods as well as conifer- grouse, blue jay, crow, wood thrush, fishes, ous and mixed forests. When feeding on abun- snakes, toads, and insects (Brown and Will, dant snowshoe hares (Lepus americanus) the 1979; de Vos 1951, 1952; Quick, 1953; fisher ranges into wetlands, alder swamps, and Hamilton and Cook, 1955; Baker, 1983). In dense coniferous undergrowth. Ordinarily, Maine (Arthur et al., 1989b) fishers ate ap- when feeding on red-backed voles, sciurids, ples, winterberry, porcupines, snowshoe and such forest prey, the fisher prefers the hares, Peromyscus, Clethrionomys, and highlands, open woodlands, cut-over forests, Microtus. Sorex and Blarina also were eat-

372 THE WILD MAMMALS OF WISCONSIN en. A few birds were also eaten, as well as are foraging for food with their mother. They deer carrion. Jon Gilbert (personal correspon- cannot kill prey until four months, and attain dence) mentioned fishers preying on Ameri- adult size at 6 months or later. Yearlings may can martens. Although habits and habitat are breed in either sex. Females will not bear correlated with certain forest prey, the fisher young until their second year. can be seen to be an opportunistic forager. Distinguishing adults from juveniles has Although the sizes of the two sexes are dis- been accomplished by determining baculum parate, the foods eaten seem to be about the development and growth of the cranial sagit- same for both sexes. tal crest (Wright and Coulter, 1967) and of Some of the remains I have observed, the suprafibular tubercle (Leach et al., 1982). confirmed and identified in Wisconsin fisher Cementum layers on the canine teeth pro- stomachs for L. Gomey and S. T. Dundei (in vide the approximate age in years. 1994) were as follows: Glaucomys sabrinus Mortality. Historically and at present 3, Odocoileus virginianus 5, Sylvilagus flori- humankind is the greatest enemy of fishers, danus 2, Sciurus sp. 4, Erethizon dorsat- by trapping and habitat destruction. Natural um 2, Tamiasciurus hudsonicus 1, Condy- enemies are unknown, but wolves could lura cristata 1, Blarina brevicauda 6, doubtless kill them, and the young and the Clethrionomys gapperi 5, Procyon lotor 1, females might be killed by some of the small- Ondatra zibethicus 2, blue jay Cyanocitta er furbearers, perhaps even by great-horned cristata 1, other birds 2. One fish and some owls (Gilbert, personal corr., Powell, 1981), plant material also were eaten. other owls, or eagles. Occasionally a fisher Reproduction. Mating takes place from kills another fisher (Jon Gilbert, personal June to September. Copulation lasts several corr.). Strickland and Douglas (1987) list pred- hours. Implantation is delayed 10 to 11 ators including, the Canada lynx, mountain months (Enders and Pearson, 1943). The lion, red fox, coyotes, eagles, great-horned mother’s embryos and the young are depen- owls, and even cannibalism. dent on her for a longer time than usual, long- Parasites on the fur include ticks, mites, er even than in Taxidea. The gestation peri- and fleas (Ixodes cookei and Oropsylla arcto- od is about 352 days. After the embryos im- mys), and internal parasites including round- plant in January, some in March or April, worms (Dracunculus, Unicinaria, Trichinel- parturition is usually in late April or May, and la, Alaria, Baylisascaris, Molineus, Capillar- occasionally as early as January. There is a ia, and others) and tapeworms (Taenia and postpartum estrus. Females breed in their Mesocestoides) (Brown and Will, 1979; de second year. The home range data suggest Vos, 1952; Dick and Leonard, 1979; Erick- promiscuity in this species. All the Wisconsin son, 1946; Hamilton and Cook, 1955; Pow- martens are believed to be induced ovulators. ell, 1981; Craig and Borecky, 1976). Strick- There is only one litter per year and it is land and Douglas 1987) list Ixodes, Listro- rather small (average 2-3, range 1-6). The phorus, Alaria, Taenia, Capillaria, Physa- young are born helpless, blind, ears closed, loptera, Uncinaria, Crenosoma, Sobolevingy- with fine hairs on the body. By 3 days the lus, Baylisascaris, Trichinella, Dracunculus, body is covered with fine grayish fur, and they the strange kidney worm Dioctophyma, and weigh about 40 g each (Coulter, 1966). The diseases toxoplasmosis and leptospirosis. eyes do not open until 50-53 days, and the Home range and density. Males have larg- deciduous teeth erupt by 85 days. By then er home ranges than females, and at least in the young are a chocolate brown color. Wean- spring overlap more than one female. There is ing occurs at 8-10 weeks, continuing until no permanent bonding of these solitary carni- approximately 100 days. By August the young vores. Not surprisingly, the males seldom over-

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 373 lap with males, and females seldom overlap with by females comprised 80 percent of theirs. Of females. Males average 20-25 km2, and females the 52 times fishers were located within an- only 15 km2 (Kelly 1977; Powell, 1981; Wright other fisher’s home range, 58 percent were and Gilbert, 1995). In Maine 7 males and 6 between sexes. Those shared areas comprised females had ranges 10. 6-78.2 km2 and 8.1 to 13 percent of home ranges of males and 69 3 9.1 km2. Movements are a few km each night. percent of the home ranges of females. Some long movements of juveniles range to 90 Of the 21 times adult fishers were locat- km in 3 days (de Vos, 1951). ed in another adult’s home range, 17 occurred In Wisconsin Kohn et al. (1993) report between males and females (four in winter, males had larger (t=2.38, P>O. 1) annual seven in spring, two in summer, four in fall). home ranges than females (15.3 vs. 3.2 mi2) , Only four instances of intrasexual overlap and the annual home range within sexes was occurred between adults. It involved two males similar for juveniles and adults. According to in spring and summer. Mean daily distance Kohn et al. (1993), home ranges of males in moved by males (1.4 miles) was greater than Wisconsin were largest during winter (8.4 mi2, by females (0. 8 mile). Fishers moved farther while those of females were largest during fall in summer than in fall or winter, and during (2.8 mi2). Home ranges of both sexes were the night than day. Snowfall and snow depths smallest during summer, probably reflecting greater than 18 inches hindered fisher move- higher food availability. Small home ranges of ments. In Maine, densities were 1 / 2.8–10.5 females during April-July probably reflected km2 in summer and 1 / 8.3-20.0 km2 in win- abundant food, the inability to travel far with a ter (Arthur et al., 1989a). litter, and reluctance to travel far from the den. Remarks. The fisher is nocturnal, although Larger home ranges of females in fall may have often active in daytime, and except when rear- reflected independence from the litter and ju- ing young or mating the animal is solitary. venile dispersal. The large fall home range of This is a tale of the alledged co-existence one juvenile female (8.3 mi2) significantly in- and possible co-evolution of two highly inter- creased the mean for all. esting animals, the browsing arboreal porcu- In 1981-82, four males shared a mean pine and the semi-arboreal fisher. In the first of 16 percent of the area of their annual home place, Seton (1953) believed the relation was ranges with other males and 17 percent with overrated, and suggested that the primary prey females. The eight radio-collared females was raccoons or mice. Powell (1979) and oth- shared a mean 15 percent of their annual ers have found that when porcupines are not home ranges with other females and 47 per- abundant fishers prey on such things as mead- cent with one or more males. Monthly home ow voles and snowshoe hares. There are few ranges for fishers were available for Novem- predators that feed on porcupines so well ber-December, 1981 and February-May, armed with dangerous quills (see account of 1982, but overlap was observed only during the porcupine), and the combat between it and the breeding season. the fisher is curious and noteworthy (Powell, Six of 12 fishers occasionally shared home 1979; Powell and Brander, 1977; de Vos, ranges in 1982-83 (Table Car-12). Juveniles 1951; Pittaway, 1978). No doubt the young were involved in 81 percent of the locations in porcupine is easily killed, but adults are a more another fisher’s home range. Most (67%) over- formidable problem. The fisher does not roll lapping of home ranges involved juvenile males. the porcupine over and attack its belly (Pow- Male home ranges overlapped 10 times as of- ell, 1981), which is devoid of quills, even ten as did those of females. The areas shared though this would seem good strategy and has by males in 1982-83 comprised 35 percent of been often reported. Instead the attack is made their annual home range, while those shared on the head and face, which are not protected

374 THE WILD MAMMALS OF WISCONSIN much by quills. The porcupine will try to hide some way. To do that would presume there its face against a tree trunk, erecting the quills are no geographic races evolved in California of the back toward the attacker. If in a tree or other places. Although there is dramatic hollow, clambering in high branches, or in a sexual variation, with males much larger than place where the head is secure, the porcupine females, and pelts that show much individual will survive, but if the fisher can open up in color and quality, there is no geographic wounds on the head and throat the porcupine variation in Wisconsin. Variation from place succumbs. The fisher rolls over the porcupine to place seems spurious or microgeographic and opens the undefended venter, ripping out (to use an old term). For Wisconsin I use the heart and flesh and leaving behind bones, in- name of the nominate race. testines and spiny skin. The fisher occasional- Specimens examined Total, 17. Iron Co.: ly picks up a few quills in its face. Although Sect. 19, 2 UW. Oneida Co.: 1UW. Lang- not an agile climber like the marten, the fisher lade Co.: Parrish Twsp, 1UW. Forest, Pine can attack a porcupine in a tree and some- counties: 12 UW (early 1900’s). Portage Co.: times forces it to descend to the ground. McDill Pond 1. Not fully appreciative of the marvelous qualities of the porcupine, the U.S. Forest Service cooperated with the Michigan and Martes americana (Turton) Wisconsin state conservation agencies to re- American Marten introduce fisher not only as a rare carnivore, but as a biological control of the porcupine. It “The fur is handsome and valuable and popularly is unlikely that fishers regulate porcupines known as American sable.” — C. B. Cory. 1912. under normal circumstances. The fisher is a spectacular hunter of the 1806. [Mustela]. americanus Turton. A general sys- snowshoe hare, following the rapid hare over tem of nature... 1: 60, type from eastern North the rough forest floor, nimbly pursuing through America. dead falls, stumps, rocks, and other obstacles, 1912. Martes americana americana: Miller. Bull. U. until it closes in for a quick attack to the head S. Nat. Mus., 79: 92. or neck, wrestling with this adversary with its 1820. Mustela leucopus Kuhl. Beitragae zur Zoolo- clawed forearms. Naturalists have often docu- gie..., p. 74. Type from Canada. mented such chases (de Vos, 1952, Seton, 1823. Mustela Huro F. Cuvier. Dictionaire des sci- 1953; Powell and Brander, 1977). ences naturelle... 29:256, type from upper Additional natural history. Powell (1981, Canada. 1982b) reviewed the biology of the fisher. 1874. [Mustela] martinus Ames. Bull. Minnesota Geographic variation. The species may Acad. Nat. Sci., 1: 69, listed with Turton’s be regarded as monotypic so long as any vari- name, referring this name to a “martin” and ation perceived is described accurately in with Turton’s name to a sable.

The name Martes means marten in Lat- Table Car-12. Mean home ranges (miles) of fishers. Moni- co Study Area. Kohn et al. 1993. in, and americana means belonging to Amer- ica. Although “pine marten” is commonly N Area Fall Winter Spring Summer used for this species, it is not restricted to pine Males 6/3.7 acres 6/8.4 4/9.7 4/2.7* forest. Therefore, some authors today call it Females 6/2.8 5/2.1 4/2.2 2/1.1 the “American marten”and some may men- Adults 6/2.2 6/3.9 6/4.6 4/2.3 tion only the word “marten.” Juveniles** 6/4.3 5/7.5 2/10 2/1.8 Description. The American or pine mar- *July only **probably young-of-the-year ten is a mink-sized slender furbearer that has

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 375 rich, shaggy fur and conspicuous ears, lined with son of total and “body” lengths, tail lengths fine buffy hairs, which held erect on the head were obtained for the following comparison: and a long neck makes this perhaps the hand- Males (N = 280) total length 584 ± 25 mm, somest mustelid in North America. Smaller and tail length 136 mm; weights 0.9 kg ± 0.1 (N = more agile than the fisher, which is described in 23); females (N = 90) 520 ± 13 mm, tail length its own account, the American marten has short- 144 mm; weights 0.6 ± 0.1 kg (N = 8). er dimensions for the teeth (upper carnassial Dental formula. I 3/3, C 1. 1, P 4/4, M labial length, less than 9.5 mm), shorter skull 1/2 = 38. (less than 90 mm), much shorter tail (less than Geographic Distribution and Status. The 300 mm), and relatively larger, more inflated American marten was eradicated but reintro- auditory bullae. From the mink (which has small- duced into Wisconsin. Two factors led to its er ears and white chin splotches) and skunks demise: loss of forests and heavy trapping for (which have black and white color patterns and its valuable sable-like fur. Formerly it was dis- flattened auditory bullae) the marten differs in tributed in northwestern Wisconsin and the having 38 teeth instead of 34. Leach (1977a, Upper Peninsula, in densely forested areas, but 1977b; and Leach and de Kleer (1978) de- was unlikely to have dwelled in grasslands in scribed the skeleton and forelimb musculature. the south and west. Today, the American mar- There are anal glands, and an abdominal gland. ten is found only in areas stocked, perhaps lost There are six inguinal mammae. The baculum even from some of them, and has not shown is described by Burt (1960), and it is tipped with the success of reintroduced fishers. There is a a perforated end. viable population ranging from northern Wis- The color is a rich yellowish brown to consin into Upper Michigan (Churchill et al., ochraceous brown, slightly darker on legs and 1981). In 2000, evidence for about 22 individ- tail, the crown brown or creamy brown. The uals were obtained (Wydeven et al., 2000). guard hairs are long and silky, but sometimes Until 1880, specimens were taken in the fur is coarse and shaggy. The eyes are Ashland County (Jackson, 1961: 331). Cory brown, and the chin spots are attractive or- (1912) in his study of Wisconsin and Illinois ange or ochraceous. The prominent ears are mammals, reported that reliable hunters and lined with buffy hairs. There is one annual trappers informed him of the marten’s sur- molt commencing in summer. vival in northern counties: Marinette, Flo- Males are larger than females, but small- rence, Price, Iron, Bayfield, Douglas, and er than the smallest fishers. Total length of Marathon. Scott (1939) mentioned one from males 600- 675 mm; tail 190- 220; hind foot 90-98; ear length 35-45; weight up to 3.2 pounds ( = 2.26 kg). Females, 540-580; tail 160-200; hind foot 78-88; ear length 34-38; weights to 2.2 pounds (= 1 kg). The skull of the American marten males averages about 85 mm in length, that of the females about 73 mm. Cranial measurements in mm of two specimens (from Forest County and an earlier record from Poynette Game Farm) in the UW-SP museum are, respectively, greatest length of skull 79.1, 76; condylobasal length 77, 74.4; maxillary tooth-row 28.1, 26.7; zygomatic breadth 44.3, 42.5; interorbital breadth 17.3, 15.9. By compari- Skull of Martes americana. E. Raymond Hall.

376 THE WILD MAMMALS OF WISCONSIN Radisson, Sawyer County, taken in l922. and Rogers, 1977; Timm, 1975) and Upper Schorger (1942: 27) reported three from Michigan (Harger and Switzenberg, 1958; Madeleine Island, in the Apostle Islands, Baker, 1983). Michigan animals came from caught by a “Frenchman” during World War Ontario, near Port Arthur. I. The last marten reported was from Maple, The American marten is beneficial to Douglas County, in the winter of 1925, but man owing to predation on rodents. Some martens supposedly persisted in Sawyer Coun- damage to songbirds is its only negative at- ty in 1939-1940 (Jackson, 1961). tribute. It seldom comes into contact with Recent reports on American martens humankind. Its lovely fur is valuable. The introduced and occurring in Wisconsin include marten does not do well on fur farms, al- those of Churchill et al. (1981) and Davis though Jackson (1961) reports young ones (1978). At an earlier date (1953) in a reintro- are easy to handle. duction now considered a failure, five mar- Habitat. The American marten occurs in tens were introduced onto Stockton Island, mature woodlands and dense conifer forest, one of the Apostle Islands in Lake Superior. ranging into open woods and meadows. Al- From 1975-1983, American martens were though this mammal reportedly prefers coni- reintroduced into the Nicolet National For- fers it certainly will range into nearby hard- est, but very few females (fewer than 27) were woods. Available foods, especially snowshoe in the 124 released. Kohn and Eckstein (1987) hares, red-backed voles, and squirrels, are determined by the 1980’s that the martens important for the American marten. See had been breeding, but almost 90 percent Mortality below. were within 12 miles of their release sites. Like the fisher the American marten These martens came from Ontario and Colo- makes use of temporary dens, but it may use rado. Since then the marten was stocked in a hollow tree or a burrow on a sunny hillside the Chequamegon National Forest. The sta- for a natal den. It often makes use of hollow tus is uncertain, except for two carcasses and trees and logs, and hides in rocky places and a few regular reports. brushpiles. In Wisconsin summer dens are The American marten has made come- almost always hollow trees (Gilbert et al., backs in Minnesota (Gunderson, 1965; Mech 1995). There is a nest of leaves and grass constructed for the young according to Jack- son (1961). In winter the marten often occupies an underground burrow. Foods. Little has been reported on foods of American martens in Wisconsin, but in other regions they feed on red squirrels, northern flying squirrels, deer mice, red-backed voles, snowshoe hares, carrion, and grouse. Mice made up two thirds of the diet, of which one quarter of the food eaten was red-backed voles. In winter the mice make up to 80 percent of the food eaten (Cowan and Mackay, 1950; Marshall, 1946). The marten is an opportunistic predator, although large mammals may not be eaten because the marten cannot kill them. Francis and Stephenson (1972), Murie Map showing most of Wisconsin in the former range of (1961), and Steventon and Major (1982) men- Martes americana in North America. tion foods: bog lemmings, birds, snakes, frogs

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 377 and toads, fishes, beetles, insects, snails, and ten) then the two similar mustelids may coex- fruits in season (blueberries, red raspberry, ser- ist. The forest type (mature for marten versus viceberry), and a little green vegetation. forest edge and cut-over woods for fisher) Reproduction. The mating act apparently might be another factor. lasts over an hour (75 min). Copulation oc- Parasites include a louse Trichodectes curs in July and August. Implantation is de- and a flea Orchopeas; these and several layed for 28-32 weeks. In February or March roundworms are the only parasites reported the process of implantation occurs and devel- by Jackson (1961). Baker (1983) lists 9 kinds opment re-commences. Parturition is about of fleas, two kinds of ticks, and four nema- 28 days later, in late March or April. The todes; probably flukes and tapeworms will be gestation period is about 220 to 275 days. found in American martens. They may suffer The young American martens are born from rabies and distemper, but these diseas- helpless, covered with sparse yellowish hairs, es are not a serious threat to martens. with eyes and ears closed. Litter size (Mead, Home Range and Density. Little is known 1994) averages two or three (range 1-5). In about the home range in this species, particu- three weeks brown hair covers the young, and larly so in Wisconsin. In Minnesota, Mech and males are larger than the females. At 26 days Rogers (1977) calculated home ranges from the ears open. Eyes open by 39 days. The young 4.2 to 8 square miles for males (= 1,088-2,122 emerge from their den at 46 days, and wean- ha), and 1. 7 square miles for females (= 440 ing takes place shortly. Young disperse as early ha). In Algonquin Provincial Park, Ontario, at 80 days, nearly full grown, and the perma- Francis and Stephenson (1972) found rather nent dentition is attained by 125 days (Brassard small home ranges: 1.4 square miles for males, and Bernard, 1939). The young may remain 0.4 square miles for females. Males wander with the mother until autumn (Strickland and more extensively than females, and probably Douglas, 1987). Aside from these observations defend territories against other males. little is known of breeding in martens. Young Additional Natural History. Strickland et American martens do not breed until their sec- al. (1982) summarized much information for ond summer (Strickland, 1975). They are aged the American marten. Jon Gilbert wrote a by counting cementum layers of canine teeth dissertation on marten and fisher ecology in and by development of the suprafibular tuber- Wisconsin. cle (Leach et al., 1982). They live as long as Geographic variation. There is no geo- 10-15 years (Crandall, 1964). graphic variation evident in Wisconsin. Mortality. The greatest enemy is human- Specimens examined. Total, 1. Forest kind. Trapping and habitat destruction are the Co: 1 mi. SE Int. Hwys. 70 & 55, Sect. 8, chief problems. Clear cutting of forests is ad- T40N, R14E.1 verse (Strickland and Douglas, 1987). Little is known about predation on martens, they are so quick and agile, and so scarce. Large Martes foina Erxleben carnivores, such as fishers (Jon Gilbert, per- Stone Marten or Beech Marten sonal corr.) and the great homed owl are suspected as natural enemies of the marten. Ei- 1777. Martes foina Erxleben. Syst. Regn. Anim. ther predation or competition for particular 1:458, type from Germany. prey may cause displacement of martens by 1792. Martes domestica Pinel. Actes Soc. H.N. Par- fishers, or vice versa (Powell and Zielinski, is, 1:55, type from France. 1983). If both prefer different foods (e.g., porcupine and raccoon for the fisher; Clethri- Description. The stone marten is a beau- onomys and squirrels for the American mar- tiful furbearer, closely resembling the rare (re-

378 THE WILD MAMMALS OF WISCONSIN Distribution. The stone marten is known from the northern Kettle Moraine State For- est, and ranges into towns and agricultural area in Waukesha, Racine and Walworth, and probably Jefferson counties (see Map). The geographic ranges are widely separated. In Europe this species ranges from Denmark and Spain to Mongolia and the Himalayas. Status. In no immediate danger of extinction in Europe, the status of the stone marten is hardly known in Wisconsin. Intro- Stone marten Martes foina.by A. N. Komarov with G. duced by a fur breeder, perhaps as early as A. Novikov. the 1940’s and more likely in 1972, it has become established in the southeastern parts introduced) American marten. The tail is elon- of the state (Long 1995). There has been no gate and bushy, the prominent ears erect (but legal protection, trapping season, or study not buffy). The limbs and tail are darker then devoted to this animal. Not common within the dorsal fur. The skull resembles that of a its new range, a few have been trapped near- mink or American marten, even a skunk. Un- ly every year, for the past 15 years. Mark like the skunk, the stone marten has a dumb- Anderson (see Long, 1995) mentioned about bell shaped upper molar and inflated auditory 20 carcasses brought in for identification in bullae, and unlike the mink (which has short, the past two years. The fur has little value. round ears) the marten has 38 total teeth, in- Occasionally this marten is raised on fur farms. cluding a full complement of premolars. The It infrequently kills chickens. ear conch is shorter and wider then those in Habitats. The stone marten occurs in the American marten. The carnassials are nar- open woods and continuous hardwood for- row and well adapted for shear. The upper molar is small, constricted anteroposteriorly, with a slightly widened internal lobe, and smaller than that of the American marten. The stone marten’s fur is less luxurious than that of the American marten; there are no orange or tawny spots on the throat. The venter is concolor with the dorsum, not creamy, buffy, or whitish as in weasels. The bold white throat pattern is striking. Total length of the stone marten varies from 620 to 840 mm, tail length from 220 to 300 mm. Weights vary from 1.1 to 2.3 kg, and the males are larger than females. Novikov (1956) records 450-540 total length, and 250-320 tail length. The condylobasal length is 76-85 mm in males, 72-78 mm in females. Zygomatic breadth varies from 46- 53 mm in males, 44-49 mm in females. Dental formula. I 3/3, C 1 /1, P 4/4, M Skull of stone marten. S. I. Ognev, G. A. Novikov. 1/2 = 38. Includes P3/ - M1/.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 379 ests, both uplands and river bottoms, and it Its nest is unknown in Wisconsin, but it in- dwells in agricultural areas, even living in hous- habits rock piles such as stone fences, and rock es and barns. Both in Europe and Wisconsin crevices, and often takes over the abandoned it inhabits farmyards and orchards. On wood- burrow of another mammal (Nowak, 1991). ed uplands and ravines forested with white Foods. The diet consists of small rodents, and bur oak, sugar maple, basswood, and such as Microtus (Hoffmeister, 1967). Birds, shagbark hickory this marten has been ob- eggs, and fruit are eaten. Vegetal matter is served and trapped in southeast Wisconsin. eaten in summer and autumn in some popu- The stone marten climbs trees with agility, lations (Novikov, 1956). In villages stone and I have a photo of one up in a tree in the martens feed on mice, rats, sparrows, what is northern Kettle Moraine Forest. They do not available, and are somewhat beneficial. climb as well as tree squirrels, but often prey In Denmark, Rasmussen and Madsen on young birds and eggs in the tree canopies. (1985) found the perecerntage of occurrence

Distribution of stone marten in Wisconsin.

380 THE WILD MAMMALS OF WISCONSIN of foods to be primarily rodents, especially ed to live along bodies of water and to swim microtines, taken mostly in rural areas, and actively. The bodies are elongate and slen- birds, mostly in urban areas. Foods were com- der, the neck especially so, and the head is pared with other regions in Europe, and al- no thicker than the neck. The brain seems to though the diet was much the same, there diminish in size with age. The dentition, al- were few lagomorphs and no lizards or shrews though resembling that of the ancient mar- in the Danish sample. Eggs of birds were eat- tens and possessing the dumbbell shaped en, and fruit was eaten in season. posterior upper molars, lost a pair of premo- Reproduction. Mating is in midsummer lars above and below. The total number of in stone martens. There is delayed implanta- teeth is 34, whereas in the more primitive tion in Europe. Total gestation varies from martens it is 38. In northern weasels, the fur 230 to 275 days. Four or five weeks of ges- molts to white (except the black hairs at the tation follow implantation. Litters are usually tail tip), and in summer weasels have brown 3-4 young, with a maximum number of 8. dorsal fur with yellowish or dirty whitish un- Lactation lasts about six weeks (Nowak, derparts. The ventral white may or may not 1991). Mead (1994) lists mean litter size as extend down the inside of the hind leg to the 2.5 (range 1-5). foot. (In the mink the color is chocolate brown Mortality. Hardly anything is known of overall, with a darkening of hairs at the tail enemies for the Wisconsin martens. Trappers tip and white splotches on the chin and kill several almost every year. No doubt domes- throat.) The fur is fine and short, often used tic dogs and automobiles will kill them on occa- by furriers for coats and decoration. The skull sion. Raptors and predators may take the young. is often said to be elongate because the brain- Home Range and Density. Nothing is case is long. The rostrum is short. The eyes known about Wisconsin stone martens. The are near the nose and mouth, and the ears home range in Alsace is about 80 ha set back. The legs in all mustelids seem rela- (Nowak, 1991). tively short for such bodies. Remarks. If the Illinois record [see records There are several interesting traits of wea- below] came from the Wisconsin population, sels and their close relatives that seem to re- then the introduction in Wisconsin did occur late to long slender bodies. The obvious bene- earlier than thought (1972). fit of being able to enter burrows of prey has Specimen examined. Northern Kettle numerous other effects (see Brown, 1972, for Moraine Forest (1 specimen mounted). the best summary), but some essential theory Other records. Photos and reports (Long, and natural history remain to be worked out. 1995). Walworth Co. Town of Troy, Lulu For one thing, any narrowing of a given body Lake; Clover Valley. Kettle Moraine State mass has a geometric effect of lengthening that Forest in Waukesha County. body, and the mathematical relations of that Illinois. Hoffmeister (1967) reported a fact have not been worked out with behavior, stone marten in northern Illinois at Blackber- physiology, mophology, and ecology of these ry Creek.

Genus Mustela Linnaeus Weasels and Mink

The genus Mustela is comprised of slender carnivores which regularly enter the burrows Sketches of heads of Mustela frenata, M. nivalis, and of mice or other small rodents, or are adapt- M. erminea. Elliot Coues, 1877.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 381 mammals. The traits needing further investigation related to the evolution of elongate, slender forms include the following: marked sexual dimorphism, with the female being much smaller; high metabolic rates but low total metabolism; short pelage; sibling competition; different diets of the sexes;”blood thirsty”behavior, and caching; territoriality and the extent of parental care; evolution of small size for carnivores; seasonality and under snow behavior; and patterns of behavior and body structure approaching convergence to small prey living in the burrows; and the absence of digging behavior, to unearth prey or excavate burrows. Note Long’s (2004) proof that elongate slenderization (prolation) increases sex divergence if each sex maintains its weight/ metabolism. Tails of M. nivalis, M. erminea, and M. frenata. The molt to white in winter seems camouflage for weasels living on snow-covered terrain, and indeed the frequency of white is high in northern latitudes. South of snow- bound regions some weasels may not turn white at all. Hall (1951) translocated weasels that molt white to warmer localities, and vice versa, and found the molt has a genetic basis.

Mustela nivalis Linnaeus Least Weasel

1766. Mustela nivalis Linnaeus. Systema nat., 12 Drawing of least weasel, winter. A. N. Komarov with ed., p.69. Type from Vesterbotten, Sweden. G. A. Novikov. Russia. 1896. Putorius rixosus Bangs. Proc. Biol. Soc. Washington, 10: 21, Type from Osler, Saskatchewan. “snow.” In Europe it is known as the mouse 1912. Mustela rixosa: Miller. Bull. U.S. Nat. Mus., weasel (= Mauswiesel). Hall (1951) recognized 79: 96. four races in North America, and there are sev- 1958. Mustela nivalis: Reichstein. Schadelvariabili- eral in Eurasia. Across the Mississippi River from tat europaischer Mausweisel (Mustela nivalis Wisconsin in Iowa is a larger, paler colored least L.) und Hermeline (Mustela erminea L.) in weasel, with longer hind foot. In Minnesota, also Beziehung zu Verbreitung und Geschlect. across the Mississippi, are the southernmost Zeitschrift fur Saugetierkunde, 22:151-182. records of the nominate race, which typically is slightly larger than M. n. allegheniensis, and The Holarctic least weasel, our smallest also has longer feet and tail (longer than 34 carnivore, derives its scientific name from Mus- mm). The Wisconsin specimens have tiny feet tela meaning “weasel” and “nivalis” meaning (up to 25 mm in males).

382 THE WILD MAMMALS OF WISCONSIN Mustela nivalis allegheniensis (Rhoads) gate and much inflated. The skull is shorter than any adult ermine skull or skull of the long- 1901. Putorius allegheniensis Rhoads. Proc. Acad. tailed weasel (basilar length less than 32 mm) Nat.Sci. Philadelphia, 52: 751, type from near and the hind foot is 25 mm or less in length. Beallsville, Washington Co., Pennsylvania. The postorbital processes flare from the skull, 1926. Mustela rixosa allegheniensis: Swenk. J. and the braincase is relatively narrower across Mamm., 7: 328. the mastoid processes and is less flattened 1983. Mustela nivalis allegheniensis: Baker. Mich- dorsally. The feet are pentadactyl with furry igan Mammals. p. 488. soles. There are two abdominal and one pec- 1959. Mustela rixosa allegheniensis: Hall and toral pair of mammae, although four abdom- Kelson. Mammals of N. Amer., p.908. Speci- inal also have been reported. Anal scent mens from Beaver Dam and Sumpter Twsp, WI. glands are in both sexes (Stubbe, 1972). Genitalia and sexual organs are described Description. The least weasel is tiny, but by Deanesly (1944), Hill (1939) and Long has typical weasel form, i.e., having long and (1969). The baculum is straight with a hook- slender body with elongate neck, flat head and like tip in adults. As in the European least large black eyes, long vibrissae, and very short weasel, M. nivalis has a karyotype of 42 chro- rostrum (and snout). The ears are rounded, mosomes with a fundamental number of 70 and the legs short, with stout feet and sharp (Mandahl and Fredga, 1980; Simonsen, claws. The least weasel is the smallest carni- 1982).). Hoffmeister (1989) lists 42 = 2N, vore in the world, about the size of a mouse with FN of 66. Latham (1953) found that the (see measurements below). The upper parts pelt or fur fluoresces curiously in ultraviolet are brown and underparts cream or white. light, not so in the other weasels. The tail is only a fourth of the head and body Upper parts are chocolate brown in sum- length, or even shorter, and it lacks a black mer, with creamy or whitish underparts. Of- tip, which is prominent in the other two Wis- ten there are a few small brown spots on the consin weasels. There are only 11 to 16 cau- venter. A few black hairs are found on the dal vertebrae. The braincase is elongate rela- short tail. In winter, except for the bright black tive to the rostrum, which is very short, re- eyes, the fur is all white. Possibly a winter sembling the slightly larger ermine’s rostrum. specimen may be found that has failed to molt, The auditory bullae are those of a hunter, elon- a phenomenon seen more often in southern latitudes, but unknown in Wisconsin. There are two molts each year, initiated by photoperiod. In spring the molt begins on the head and back and works posteriorly and ventrally. In autumn, the reverse occurs (see account of the long-tailed weasel below). In Wisconsin white is attained in October and November, and brown fur is attained in spring, usually by early May or sooner. Specimens from Iowa County and Monroe County (21 March) are changing to brown. White is a concealment color matching snowy surroundings. The reduction (suggested by the few black hairs present) of black in the tail results obviously from natural selection at work, because the tail is so short, if Skull of Mustela nivalis. it were conspicuous, it would draw the strike of

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 383 a raptor to the body. In larger weasels the black tail tip draws the strike away from the body (see Mustela frenata below). The hind foot is less than 27 mm, tail shorter than 35 mm, basilar length of skull shorter than 33 mm. Males are perhaps 10 percent longer than females in measurements, and heavier (up to 60 percent, perhaps 100 percent heavier than females). The males range up to 206 mm in total length, to 40 mm in tail length, usually to 23 mm in hind foot length, and to 21 mm in length of ear. Weights are reported up to 60 g in males.

Maps showing geographic distribution of Mustela nivalis in Wiscsonsin and North America.

384 THE WILD MAMMALS OF WISCONSIN Three skulls from central Wisconsin are latest records are northern extensions of short, M 33.2 mm, M 31.5, F 24; cranial range. If Hoffmeister (1989), following Hall, breadth 14-15. is correct in assigning Illinois specimens to Hanson (1945) provided measurements M. n. allegheniensis, northern range exten- for six least weasels from Sauk and Dane sions of a prairie species are more likely to counties, March-April, 1943, excepting one explain the occurrence in Wisconsin than male taken in November, as follows: One emigration eastward across the Mississippi March male weighed 60 g, and a female 48.3 River, and across the several other wide riv- g. Total length varied from 182-198 mm in ers in western Wisconsin. The possibility that all the males, 165-184 in the females. Tail some immigration from the west occurred length varied from 26-34 mm in the males, must be recognized as a possibility; it hap- 23-27 in the females. Hind foot varied from pened for Spilogale, Geomys, and Pitymys. 22-24 mm in the males, 19.5-22 in females. Status. Although I hear of recent occur- Some Wisconsin weasels are so difficult rences of least weasels in Walworth and Ra- to identify in the flesh, either in winter or sum- cine counties, there has been only one taken mer pelages, characters are mentioned here in central Wisconsin in the last ten years. All that usually help identify the adults. In size the weasels are uncommon, except for local the small weasels are M. nivalis or female M. occurrences of ermines (M. erminea). The least erminea (which always have black-tipped weasel is the rarest, and the recent prohibition tails). If the specimen is of medium size, males of some insecticides and protection of wetlands are usually ermines and the females are usu- may increase the low densities of M. nivalis. ally M. frenata (which usually have a tail length The least weasel needs management as greater than 40-45% of the head and body well as protection. Weasel trapping might be length). prohibited in some areas, and the species’ hab- Dental formula. I 3/3, C 1/1, P 3/3, M itat requirements need to be ascertained. Its 1/2 = 34. only fault might be the killing of an occasion- Geographic Distribution. This species is al snake or small bird, and its benefits to hu- circumboreal (Allen, 1933) but in Wisconsin mankind in controlling rodents far exceed any has always been rare, and in the boreal north- possible detriment. Criddle (1926) mentioned ern counties, seems extremely rare. It would how least weasels cleaned out an area inhab- seem on geographic grounds that the Wis- ited by 27 groups of meadow mice (average consin least weasels would belong to a near- group size 4.5 voles), using the burrows of by western race rather than to a Pennsyvania the mice from which to strike out for new race. Although the geographic variation in the attacks. Its fur is not valuable. three races is slight, there was no taxonomist Habitat. Mustela nivalis is most com- more careful than E. R. Hall (1951) in mea- mon in marsh lands, fields, and wet prairies, suring skulls and appraising variation. The ranging into towns and fields on occasion, in least weasel is a prairie species ranging into the southern and western counties. On the Wisconsin northward from the prairie penin- Michigan State University campus along rail- sula of Illinois. From southern localities it road rights-of-way, Gary Heidt caught them seems to be extending its range northward, frequently in mixed grasses, weeds and shrubs following eastern farm lands and cleared fields (Baker, 1983). The least weasel also prefers from marsh to marsh even onto the Upper woodland edge, railroad right-of-way, well- Peninsula of Michigan. A pincer of sorts re- drained prairies, grassy fields, old fields and sults from this extension of range and also a grain fields, and brushy fencerows. For the western invasion (1975) into Upper Michigan most part these habitats are found intermit- (Baker, 1983). The map shows several of the tently in the heavily farmed eastern counties,

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 385 the western counties, and of course along the implantation. Heidt (1970) and Heidt et al. southern border of Wisconsin. (1968) have summarized reproductive behav- Ermines, which are only a little larger ior in this species. than least weasels, compete with least wea- Gestation lasts about 34 to 37 days re- sels but when food is abundant they coexist, sulting in a litter of from one to six young. fluctuating in numbers depending on environ- Mating takes place in any month of the year, mental factors (Powell and Zielinski, 1983). even winter, but spring and autumn seem They probably feed on different foods in dif- more favorable. There are probably two lit- ferent habitats when prey is in short supply ters, possibly three per year if food abundance (Simms, 1979). Coexistence is only tempo- permits. There is transuterine migration of rary, resulting in the disappearance of one blastocysts to insure proper distribution of and possible re-colonization at a later date. fetuses. The young are born naked, pink and The nest is in a shallow burrow made by blind. They crawl about using their forelimbs. a mouse or other rodent. Ellison (1946) found In a week a coat of fine whitish hair covers a least weasel within the burrow of a mole. the body, and the eyes open and teeth erupt These dens are usually in meadows or fields. by 11 days. By 18 days the coat pattern is Jackson (1961) reports one from a corn shock evident, the young are almost furred out, and in Wisconsin. The nest is made of grass or they attempt to eat meat. After birth the young twigs lined with fur from the mice eaten. This make high squeaks, replaced by chirps after matted nest fur may be an inch thick. Other the eyes open. Trills do not appear until after nest sites in the burrow are utilized, but the 35-50 days (Heidt and Huff, 1970). By 40 weasel sleeps and rears its young in the furry days the young can kill mice, and by 95 days nests. Partly eaten carcasses are often found they attain adult weight. Females are repro- in the burrow, and feces and bones are strewn ductively viable after only four months, and about outside. Criddle (1947) also described the male breeds after eight months. Aging of a least weasel’s nest. Several dens are used in least weasels has been attempted by using an animal’s home range. wear on the carnassial and counting dental Foods. Foods are usually discussed in re- annuli. Neither has been related to actual age gard to European least weasels, but certainly of least weasels. the least weasel is a predator specialized to en- Mortality. Almost any raptor, large snake, ter the small burrows of mice. The smaller fe- or carnivore may kill these little weasels, even male spends more time preying in burrows than domestic cats and dogs. Humans kill them in the larger male, which feeds more often on birds, traps and with automobiles. eggs, insects, and young rabbits. The diet is that Hardly anything is known about preda- of an opportunist, but the staple food, and the tors or parasites in North American least wea- food impacting territory and breeding area, is sels. Skrjabingylus nasicola infects the nasal the abundance of mice. Even in winter the pre- sinuses, and other worms (Taenia, Physa- dation continues on and under the snow. loptera, and Moliens) are internal parasites. Maintenance requirements for captive M. Doubtless fleas, mites, and ticks are found on nivalis are about 1 g food (wet weight) per them, as in Europe. hour. A 50-60 g (1.75-2.1 oz) weasel eats Home range and density. Not surprising- 40-60 per cent of its own weight daily (Gill- ly, there is little information on home range ingham, 1984). In practical terms this equals and density of least weasels in Wisconsin. one or two mice per day. Reportedly males wander much farther than Reproduction. Mating has been de- females, both circumscribed to territories (ex- scribed by Sheffield and King (1994). Copu- cept in the mating season when the sexes keep lation induces ovulation. There is no delayed separate) related to fluctuating food supply.

386 THE WILD MAMMALS OF WISCONSIN In England, King (1975) found home range defend their dens against males when in late of males ranging from 17.5-37.5 acres (7-15 pregnancy and during lactation. Mating is ha); that of females no more than 10 acres (4 promiscuous, and males do not help rear the ha). In this country perhaps 2 acres is a fair young. Vocalizations include chirps, hisses, estimate (Baker, 1983). Home range in this trills, squeals, and squeaks. small predator must include subterranean Additional natural history. Biology of wanderings in the burrows of mice. These the least weasel is reviewed by Sheffield and extended home range areas are an enigma. King (1994). Hanson (1945) caught five in prairie and old Geographic variation. Although least field habitats in eastern Sauk County, Wis- weasels along the Mississippi resemble wea- consin, in the spring of 1943. sels of the subspecies westward, in Iowa or Remarks. As in other weasels the hunt- Minnesota, there is no evidence of geographic ing is by sight, smell, and hearing, as the variation in Wisconsin. weasel moves quickly and “tirelessly,” search- Specimens examined. Total, 30. Dane ing holes and crevices along regular hunting Co.: 13 UW, 1 UW Wildl. Ecol. Dunn Co.: routes. These routes are systematically Lawler Rd.1 UW. Grant Co.: 4 mi. S Lan- changed, rotating from one part of the habi- caster 1 UW Wildl. Ecol. Iowa Co.: Mineral tat to another. Frequently the weasel stands Point 1UW. Lafayette Co.: Fayette Twsp on its hind feet looking about, and I had one 1UW. Manitowoc Co.: Manitowoc, Sect. 18, captive for a few days that looked up at me, 1. Marathon Co.: Wausau 1 UW. Monroe to my thinking, as possible prey. That makes Co.: Sparta 1 UW. Portage Co.: Stevens more feasible the report in Ognev (1931, Point, 1. 15 mi. E Stevens Point 1. Buena 1962) that a European least weasel was found Vista Marsh 1. Racine Co.: Hwy 38 and 31, repeatedly (for several nights) riding on the 1. Vernon Co.: Viroqua 2. Washington Co.: neck, entangling the mane of a frantic, sta- 5 mi. E West Bend 1. Waushara Co.: T20N, bled horse, which showed tiny blood spots R8E, Sect. 8, 1. on the neck. The least weasel bounds along in one- to three-foot bounds, and in Europe is known Mustela erminea Linnaeus as an able swimmer (Sheffield and King, Ermine 1994). This hunter is active day or night, any month of the year.. Activity depends on the 1758. Mustela erminea Linnaeus. Systema naturae, abundance and activity of the prey (Zielinski, 10th ed. 1988). Killing is not learned; even captive, 1838. Mustela cigognanii [sic. = cicognanii] Bona- motherless weasels kill by nature. The “blood- parate. Charlesworth mag. Nat. Hist., 2:37, thirsty” nature of weasels is legendary, and type from eastern United States. weasels kill until they are too tired to contin- 1838. Mustela richardsoni Bonaparte. Charlesworth ue (Heidt, 1972). In the den the young wea- Mag. Nat. Hist., 2: 38, type probably from Fort sels wrestle and play. Except when breeding Franklin, Canada. these weasels are solitary. 1896. Putorius streatori Merriam. N. Amer. Fau- Killing of mice is accomplished by an at- na, 11: 13, type from Mount Vernon, Wash- tack on the nape (or back) of the skull, or the ington. weasel tears open the victim’s throat. The 1896. Putorius arcticus Merriam. N. Amer. Fauna, brain is preferred as food. Caching is ordi- 11: 15, type from Point Barrow, Alaska. nary and perhaps necessary for this tiny, high- 1899. Putorius (Arctogale) muricus Bangs. Proc. ly active predator. The den must be defend- New England Zool. Club, 1: 71, type from ed, except from family members. Females Echo, El Dorado, California.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 387 Mustela erminea bangsi Hall fantile hairs to adult pelage. The brown color varies from place to place, coffee brown, milk- 1945. Mustela erminea bangsi Hall. J. Mammalo- coffee brown, rich chocolate brown, and gy, 26: 176, type from Elk River, Sherburne sometimes grayish brown. In molt the mix- Co., Minnesota. ture of color seems muddied and whitish brown. In winter the white fur may be tinged The name Mustela, from Linnaeus, or washed with a bright canary yellow tint. means weasel in Latin. The name erminea Males range from 225-340 mm, females means white winter coat, indirectly referring 190-290, with the tail only 30 to 45 percent to an Anglo-Saxon deity Eormen (Baker, of the head and body length. The tail verte- 1983). Used in the vernacular the word er- brae number only 16-19. The postglenoid/ mine means white weasel fur. In Europe the condydlobasal ratio is large, exceeding 48 in word is Hermelin or Hermine (silent H), females and 46 in males. In long-tailed wea- where once I saw a cape belonging to Kath- sels it is less. This ratio reflects the short ros- ryn the Great containing, I estimate, some- trum. See Table Car-13. what more than 300 ermines with their little black-tipped tails attached. The name bangsi honors a turn-of-the-20th century New En- gland naturalist O. Bangs, who named several American weasels (see Synonymy above). Description. The ermine is a small weasel (see measurements below), the females often as small as least weasels and the males approximately as large as female long-tailed weasels. The tail is short (less than 40 per cent of head and body length), distinctly pencilled with a short tip of black. The rostrum (snout) is short. In summer fur the feet are often brown with an inner line of whitish, although there are Figure on characters of weasel skulls. Dorsal aspects of some ermines with all brown legs. In addition the skull in ermine (small) and long-tailed weasel. Note shape of the rostrum and postorbital processes. to a short, arcuate rostrum, the ermine has no distinct, flaring, spear-pointed postorbital processes. They are pointed protruberances, but not so triangular. Other comparisons are in the accounts of M. nivalis and M. frenata. The baculum was described by Burt (1960) and the penis by Long (1969). The ten mammae are inguinal and abdominal. The karyotype is 2N = 44 (King, 1983). The ermine is brown above in summer and whitish or buff below, with black pencilled tail and black eyes. In winter probably all Wisconsin ermines molt to white. This molt commences in late October and early Novem- ber. There are two annual molts induced by photoperiod (Hall, 1951, and others). There also is a molt in the juvenile from whitish in- Skull of Mustela erminea.

388 THE WILD MAMMALS OF WISCONSIN For diagnosing ermines from other weasels in Wisconsin, see their accounts, see Mus- telinae, and figures on weasel skulls and tails. Dental Formula. I 3/3, C 1/1, P 3/3, M 1/2 = 34. Geographic Distribution. The ermine is unknown in the southernmost counties, although a record from Waukesha County in the east, and another from northeastern Iowa across the Mississippi near Galena, Illinois, suggest that on occasion an ermine crosses the border into Illinois, where it remains unknown. In southern Wisconsin there are few

Maps showing geographic distribution of Mustela erminea in Wisconsin and North America.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 389 boreal habitats and possible competition with a microtine vole. Nests are often in piles of least weasels and long-tailed weasels. The rocks in Eurasia (Ognev, 1931, 1962) and absence of deep snow and the presence of eastern America. The nest is usually a mass other carnivores, taken together, may exclude of dry vegetation often lined with fur or feath- M. erminea. Ermines are widely distributed ers of victims, with side cavities off the bur- in northern and central Wisconsin. There are row utilized for carcasses and latrines. Hol- specimens in this collection from the Apostle low logs, woodpiles, and stone walls also are Islands (see below). used for homes. Winter dens are often sur- Status. Except in the southern and west- face nests of Microtus (e.g., Fitzgerald, 1977). ern counties, and in urbanized areas, the er- Foods. Although the ermine is sexually mine is fairly common for a predator, and is dimorphic, the foods of the two sexes are not the most common Wisconsin weasel. It is de- much different, consisting of the most abun- cidedly beneficial, an efficient mouser, and dant rodents for the most part (Simms, seldom is any problem to humankind. It 1979a). Some carcasses are cached. About wreaks havoc by surplus killing if it enters a half the diet is mice, but shrews are occasion- chicken yard. Fortunately, people and ermines ally eaten, and hares and rabbits are often are not often in contact. The winter fur has killed for food. Audubon and Bachman (1842) some value, more so in the past. The aes- mention an entire family of chipmunks killed thetic qualities are valuable, for sight of an by an ermine, and stated that an ermine can ermine in the wild is a life-long experience. kill a rabbit (Sylvilagus) or grouse (Bonasa There are some areas where local protection umbellus) ten times its own weight. One re- of ermines is worthwhile, particularly along leased under a barn floor killed and dragged the southern margin of known range. together a pile of rats (Rattus norvegicus). Habitat. The ermine is generally confined The ermine feeds opportunistically on vari- to boreal habitats such as marshes, northern ous rodents, birds, reptiles, amphibians, fish- wet sedge meadows, a variety of old fields, es, earthworms, and carrion (Aldous and meadows, clearings, and even beaches. Er- Manweiler, 1942). mines are found in a great circumpolar distri- Reproduction. Breeding commences in bution, and Baker (1983) suggests the lack of early summer, including the young nestling continuous snow cover in Indiana, southern females. Copulation lasts 2-20 minutes and Michigan, southern Wisconsin, and northern may be repeated. The pair bond is not main- Illinois might contribute to scarcity or absence tained thereafter. Implantation is delayed. The of ermines in the south. He also suggests the delay in implantation (10-12 months) and possibility of competition between ermines gestation (28 days) are more than 300 days. and the southern weasels. The ermine is of- In March the embryo implants, and parturi- ten found in successional forests, seldom in tion occurs in mid-April or early May (Simms, heavy forest (which is the domaine of the 1979a; Wright 1942; Fagerstone, 1987). martens), and dwells in brushy grasslands near About six embryos (1-9) are born, the young swamps, marshes, and streams. The species nearly naked or covered already with fine may occur at the forest edge, near bogs, and white hair. They weigh as much as 3.5 g each in dry hardwoods. An introduced pest in New (usually about 1.7 g) and their eyes are closed. Zealand, it regularly climbs trees in the for- A brownish mane appears by 2 weeks. After ests to enter the nestholes of birds. This be- a week they are furred out and make squeaks. havior is not general for weasels in America By three weeks the deciduous teeth are erupt- (but see Rutter, 1930; Booth, 1946). ed, and the young weighs about 16 to 17 g As in other weasels, the nest is taken over (5.6-6 oz). At 35 days the eyes open and the from another animal, usually a rodent, often young show the summer color pattern. They

390 THE WILD MAMMALS OF WISCONSIN eat meat provided by the mother. By 45 days ticks, and lice infest the fur (Jackson, 1961; the young wrestle and play, follow their moth- Scharf and Stewart, 1980; Wilson and er about, and are weaned soon after. The Johnson, 1971; Lawrence et al., 1965; Fag- young females themselves may breed when erstone, 1987). Diseases include tularemia 60 to 70 days of age. Ermines live in nature and distemper. only about two years or even less. Home range and Density. Males have Two small ermines from a litter of 7 tak- larger home ranges, overlapping female en 14 May 1956, at 1.5 m. E Prior Lake in home ranges, and sometimes using home Scott County, Minnesota, showed replacement areas of two females. Males do breed with of the fine hair with fur on the crown and nape. more than one female, and both sexes tend These measured 123-21-17 (female) and 132- to defend their home areas against juveniles 24-19 (male) and weighed, respectively, 13.4 and other adults (Powell, 1979). All the wea- g (0.46 oz) and 19.0 g (0.67 oz). sels tend to be solitary except when breed- Mortality. Raptors and carnivores often ing or females maintaining families. The av- capture ermines, in spite of their quick move- erage home range was 63.2 acres (25.3 ha) ments and fierce nature. Their small size leads for males, 29.2 (11.7 ha) for females (Simms, to such predation. Enemies include coyote, 1979). One movement was 1,800 feet (549 red and gray foxes, marten, fisher, badger, m) in 24 hours, and males were more active house cat, black snake, doubtless the fox travelers than females. snake, rough-legged hawk and goshawk in Simms (1979) found in Ontario the av- winter, great horned owl, barred owl, snowy erage monthly density was one ermine per owl, and bald eagle (Hamilton, 1933; Ban- 278 acres, but if he counted only the succes- field, 1974). sional habitats the number increased to one Parasites include Alaria mustelae, Alar- per 160 acres area (64 ha). Densities varied ia taxidae, Taenia, Dracunculus, Physa- seasonally, highest in summer as young en- loptera, and Molineus, the nasal parasite tered the population. Skrjabingylus nasicola, and Moniliformis Remarks. Audubon and Bachman (1842) (Jackson, 1961; Baker, 1983). Fleas, mites, mention the adaptation to snow, that the ermine travels by long leaps over the snow, and makes “galleries” or tunnels in deep snow, Table Car-13Car-13. Cranial measurements of adult ermines by some extending 20 to 30 yards. The family sex. Records include specimens from Field Mus., Univ. Mich- stays together including mother and young, igan, Michigan State, and UW-SP.* even the males grown large until autumn. The young seldom leave the natal “locality.” Gen- Locality Total Tail Hind Cond. Cranial Max. l. foot l. br. t-r erally considered a fearless and vicious animal, Leonardo da Vinci painted an ermine Females being held by a woman, suggesting it was a Upper Mich 234 45 31 35.0 15.5 9.4 Upper Mich — — — 35.7 16.1 9.5 docile pet. Forest Co 260 60 34 36.3 15.3 9.7 Additional Natural History. Carolyn King Price Co 259 58 30 36.3 17.5 10.2 (1983) wrote a review of ermine biology. Males Geographic Variation. There is only one Rusk Co 301 71 44 42.7 18.4 12.1 geographic race in Wisconsin. Portage Co 303 81 44 43.4 19.6 12.3 Portage Co 244 67 35 39.2 18.1 10.7 Specimens examained. Total, 58. Ash- Portage Co 260 72 35 39.0 19.0 10.6 land (Madeleine Island 1 UW. Outer Island, Dodge Co** 235 60 30 36.1 17.8 9.7 Apostle Islands 2. Butternut 1 FM. 18 mi. W * Ratios of tail/head & body < 39, N = 16 Butternut at Bear Lake 1 FM. Bear Lake 2 **Age subadult UW), Barren, Calument, Clark, Columbia:

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 391 Wyocena Twsp 1. Dodge Co.: Horicon N. W. R., 1 Field Mus. Door Co.: Hwy 5, 2 mi. S Sturgeon Bay 1. 6 mi. W Sturgeon Bay 1. Fish Creek 1 UW. Florence, Fond du Lac, Forest, Green Lake.: Berlin 1. Iron, Jack- son, Juneau, Lincoln, Manitowoc, Mara- thon, Menominee, Oneida, Outagamie, Oza- Sketch of long-tailed weasel. V. Hogg, in Hall. ukee Co.: Cedar Grove 1. Portage, Price, Rusk, Sheboygan, Waupaca, Waushara, Wood counties. The name Mustela means weasel in Lat- Michigan. Delta Co.: 1/2 mile S, 1 1/2 in, and frenata means having a “bridle.” In mi. E Gorden 1 MSU. Gogebic Co.: Ironwood the southern parts of the range, this species 1 UMich. Cisco Lake 1 UMich. Iron Co. 7 mi. often has a mask-like “bridle” on the face. S Crystal Falls. Minnesota. Scott Co. 2 The name was originally applied to “bridled” weasels from Mexico. The subspecific name noveboracensis means “from New York.” Mustela frenata Lichtenstein (See account of Peromyscus leucopus noveb- Long-tailed Weasel oracensis.) Description. The male longtail is a large 1831. Mustela frenata Lichtenstein. Darstellung nen- weasel, and the female may be medium or er oder wenig bekannter Saugethiere..., pl. 42 small. Some are smaller than female ermines. and unpaged text. Type from Ciudad Mexico, The body is the typical weasel shape, long Mexico. slender body and short legs, pentadactyl and 1877. Putorius mexicanus Coues. Fur-bearing ani- clawed feet, an elongate neck and narrow mals. U.S. Geol. Surv. Territories, a nomen head, with bright black eyes and erect round- nudem, cited as a name used by Berlandier ed ears, the head held high like a serpent, from Tamaulipas and Metamoras, Mexico. and having a long black-tipped tail some 40- 70 percent of the length of head and body. This slender carnivore is adapted to entering Mustela frenata noveboracensis (Emmons) the narrow passages and runways of mice. The skull has a longer rostrum (snout) than 1840. Putorius noveboracensis Emmons. A report the other weasels, resembling that of a mink on the quadrupeds of Massachusetts, p. 45. N. vison. The longer rostrum causes the post- Type from Williamstown, Berkshire Co., Mas- glenoid/condylobasal ratio to be less than 47 sachusetts. percent in females, 46 in males. The postgle- 1936. Mustela frenata noveboracensis: Hall. Carn- noid length is the distance between the gle- egie Inst. Washington, Publ. 473: 104. noid fossa or jaw socket and the posterior- 1842. Mustela fusca Audubon and Bachman. J. most point of the exoccipital condyle. The Acad. Nat. Sci. Philadelphia. 8: 288, type from postorbital processes, in fully adult weasels, New York. stand away from the skull and are triangular 1853. Putorius agilis Audubon and Bachman. The or pointed. There are 19-23 caudal vertebrae. viviparous quadrupeds of North America. 3: Often identification is difficult because an 184, pl. 140, type from New York. Not the occasional M. frenata has a short tail, resem- Peruvian weasel Mustela agilis Tschudi, 1844. bling M. erminea. Many weasel skulls are 1899. Putorius noveboracensis notius Bangs. Proc. crushed on roads. There is overlap in size with New England Zool. Club, 1: 53, type from ermines. Some ermines have somewhat point- Weaverville, Buncombe Co., North Carolina. ed postorbital processes, some longtails are

392 THE WILD MAMMALS OF WISCONSIN runts, even adult males, and always we have melanin pigments in the fur) when transplant- the problem identifying young animals. There ed into warm areas; those that kept the brown is much individual variation in specimens from color in winter did so when transplanted into southern Wisconsin and Illinois, so that some cold habitats. This suggests a genetic basis. of the specimens there do not much resem- The summer pelage on the dorsum con- ble others. The aforementioned characters will sists of long guard hairs of cinnamon brown work most of the time. Additonally there are and dense grayish-tawny underfur. Molt lasts the Measurements of tail and hind foot (see about 25 days, in the period from February Table Car-14). The black tail pencil is longer to April, to attain brown upper parts, and and less set off from the brown or white fur from October until late November to attain in frenata (see figure on tails). white winter fur. Fur is mottled and pale There are four abdominal and four in- muddy brown at these times. Photoperiod guinal mammae. The penis was described by initiates the molts, and the white provides Long (1959), and the baculum figured by Burt concealment in snow. The long black pencil (1960). The chromosomes number 2N = 42, draws the strike of the raptor away from the with a FN of 66 (Sheffield and Thomas, 1997). body of the weasel, and is obviously adap- In Wisconsin the long-tailed weasel is tive. Experimental evidence by use of a pet white in winter, excepting its black eyes and owl and pet hawk confirmed this hypothesis black pencilled tail. The winter fur may be by allowing the birds to attack moving stuffed stained, probably by urine and musk, to a weasels on wheels (Powell, 1982). Perhaps bright lemon yellow. In summer, the color is to match the long-lasting snow cover in coffee brown above, creamy yellow or whit- northern lataitudes in winter, while avoiding ish below. The ventral pelage in summer is the contrast of white upper parts in other often tinged with orange, but the reportaed seasons, the two-toned long-tailed weasel absence of an inner line of creamy buff (con- delays molt to white by progressing from color with underparts) extending down the leg venter to dorsum in fall, and rids its upper to the hind foot is not a good character (some- parts of white quickly by progressing the times present in M. frenata, sometimes the opposite in spring. line is lacking in M. erminea). The sexes are greatly dimorphic, with the Hall (1951) reported weasels that were males usually much larger than the females. white in winter kept their color (absence of The males are at least 10 percent larger in

Skull of Mustela frenata. Also see sketch of weasel skulls. Long-tailed weasels in white and summer brown pelages. Note postorbital processes. G. Heidt.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 393 external measurements, except in a few runty males; and the weight may be as much as double that of the female. Greatest length of the skull in males is 44-51 mm, in females 37-45 mm. External and cranial measurements are given in Table Car-14. The tail is usually longer that 110 mm (4.3 in) in males, more than 80 mm (3.12 in) in females, and the hind foot is usually 45 mm (1.75 in) in males, 35 mm (1.36 in) in females. Dental formula. I 3/3, C 1/1, P. 3/3, M 1/2 = 34.

Maps showing geographic distribution of Mustela frenata in Wisconsin and North America.

394 THE WILD MAMMALS OF WISCONSIN Geographic Distribution. The Mississip- of seeing one in nature is an asset seldom pi River bounds the range of this race of long- considered, and seldom enjoyed. I have seen tailed weasel (Hall, 1951). Westward in Min- only one of them alive in the wild, and for a nesota is the race M. f. spadix, and in Iowa, very brief period of time running along a road- ranging up to the Wisconsin border, is the side ditch and up the embankment. race M. f. primulina. The weasel M. f. novab- Habitat. The long-tailed weasel is found oracensis is found throughout Wisconsin in in clearings of forests and old fields, wood- suitable habitats, also throughout Upper and lots, the edge of marshes, hedgerows, barn- Lower Michigan, and southward in Illinois. yards, and brushy grasslands, often near Status. Of the three weasels in Wiscon- creeks or lakes, and even on sand dunes. sin, this species is second in abundance, and Home ranges are associated with high popu- first in the southern counties where the er- lations of mice. Longtails have been reported mine is unknown and the least weasel rare. It from swampy meadows with aspens, along prefers cat-tail marshes and dense cover (Shef- river courses, and other areas where rodents field 1997). It is found in wetlands with either are abundant. They often dwell in wetlands, of the other weasels on occasion, but often or near standing water. inhabits higher, dryer habitats than they. How this habitat compares with that of the Unlike the boreal ermine (M. erminea) the ermine is unclear, being quite similar, but per- long-tailed weasel ranges southward through- haps it is somewhat more open (i.e., vegetation out a vast region in temperate and tropical less dense). Geographically wide-spread, the America. Probably an early invader into Wis- long-tailed weasel has no close relative in Eur- consin from the south, it followed the retreat- asia, as do the two other weasels, and seems to ing boreal habitats, perhaps, occupying the have ranged from the south and east northward, uncovered habitats after the Pleistocene (Wis- reaching limits in the snowy parts of Canada. It consinan?) glaciation. shows close affinity, i.e., relationship, to the The long-tailed weasel seems less adverse- South American Mustela africana, which is ly affected by land use than the other weasels. more primitive. Occurring sympatrically with From the tip of the Door Peninsula and from ermines in western states and provinces, the widely distributed records on the Upper Pen- long-tailed weasel is absent from deserts. Stand- insula, the long-tailed weasel seems to be ex- ing water, mentioned as important, is abundant cluded from suitable habitats only by water in Wisconsin. Food availability, i.e., rodents of barriers, from the Apostle Islands in Lake Su- various kinds in open areas, and shrubby cover perior, and the isles that traverse Green Bay where birds and rabbits can be hunted, seem in Lake Michigan. To humankind this weasel the most important factors (see Fagerstone, seems more a pest than are the other two, 1987). Decayed logs and fallen trees for natal although it kills many mice and enters barns to dens seems important. kill mice and Norway rats. It also frequently A den is partially described by Kennicott kills cottontails, which to many people is pre- (see Foods below). Ingles (1939) also described dation apparently beneficial to humankind. The the nest. Usually the weasel takes over a bur- longtail may become a chicken killer. row of a chipmunk or ground squirrel, or some The winter pelt is more valuable than that other rodent. Weasels seldom dig. According of the ermine. Neither fur is valuable at the to Audubon and Bachman (1842) they are present time, because animal rights groups have quite clumsy at it. Bodies of mice and partially depressed the fur market. Also the pelt is small. eaten carcasses, even of birds, are often found Jackson (1961) considers this weasel to in the appropriated burrows (Sheffield and be more of an asset than a problem, chicken Thomas, 1997; Seton, 1953). Hall (1951) re- killing notwithstanding. The aesthetic value viewed descriptions of dens of the long-tailed

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 395 weasel from its wide range, and some data on sin farmer who observed a female long-tailed natal dens are listed here: Female and 8 young, weasel carrying two mice into a log, which he burrow 3 in diameter, two chambers depth 12 sawed open finding five young weasels and a in, length 3 ft (1 m), nest material chopped large number of cached mice. The mother sub- grass; Female and 4 young, diameter?, length sequently moved the young to a more secure 10 feet in a hollowed, rotted fallen oak and place. Sheffield (1997) provides a food list of adjacent pocket gopher burrow, nest materi- a varied regimen, including these Wisconsin al?; Mother and 3 young, “a bushel” of rabbit, kinds: Microtus, Clethrionomys, Peromyscus, rat and squirrel hair, dimensions?; Female and Synaptomys, Ondatra, Zapus, Mus, Rattus, 3 young, hollow pine log, dimensions and Geomys, Spermophilus, Tamias, Eutamias, material?; Mother and 3 young, rodent bur- Tamiasciurus, Sciurus, Glaucomys, Sylvila- row, dimensions and material?; Female and 4 gus, Lepus americanus, Scalopus, Condylu- young, beneath decayed log, nest of Microtus ra, Sorex, Microsorex, Blarina, Eptesicus, fur, dimensions?. Other nests described were birds such as Agelaius, Spizella, Colinus, and a bag of feathers; a bunch of straw under a others. The duck, Anas discors, some snakes threshing machine; a mole burrow with nest and lizards, other weasels, and numerous in- of grass (9 by 6 in) with internal nest-hollow sects also were eaten. and patches of mouse and shrew skins; a Reproduction. This weasel is generally ground squirrel burrow with nest 7 by 9 in, solitary except when mating and rearing young. with some mouse and rabbit fur and shrew Unlike the least weasel, the long-tailed weasel remains; and a ground squirrel burrow with has a long delay prior to implanting the blasto- grass nest 11 by 9 in, depth below ground sur- cyst embryos. Wright (1948) described the preface 11 in, with remains of mice, near an aban- implanted blastocysts. The gestation is about doned skunk’s nest. 270 days, and after the blastocysts are implant- Foods. The chief foods of Mustela freed parturition follows about 21 to 28 days lat- nata include mice, especially meadow voles, er. The young are probably born in April in cottontails, and birds (especially ground nest- Wisconsin, and emerge from the den in June. ers and feeders, including domestic chickens), Embryos range from 1-9, perhaps 10, but the and also worms (Osgood, 1936: 64), lizards, usual number is 5 or 6. The young are born and snakes. Males often take larger prey than pink and almost naked. The eyes open by the females. In Iowa Polderboer et al. (1941) 37th day. Weaning is not much later than this. found in scats the following foods and fre- In 12 weeks the permanent teeth and mature quencies of capture: size are attained. Males cannot breed their first Number Kind Frequency year, but females may, and usually breed their 71 Microtus 43% second year and thereafter. Age groups have (includes some I regard as Pitymys) been set forth by Hall (1951) as follows: Juve- 36 Reithrodontomys 22% nile- deciduous teeth, birth to 3 months; 17 Peromyscus sp. 10% Young- suture widely open between maxillae 14 Sylvilagus 8% and nasals, and between premaxillaries and Number, Kinds, Frequency nasals 3-7.5 months; Subadult- aforemen- 22 Other mice, Geomys, tioned sutures indistinct but visible 7.7-10 and least weasel 13% months; Adult- sutures closed 10 months or 1 Tree sparrow 0.6% (?winter) older. The young have a crest of hair, called 1 Grasshopper 0.6% (?summer) the “pompadour” on their heads. Both sexes ?1.85% seem to protect and attend the young, and These foods were studied for winter and bring prey to the den (even worms, Hamilton, spring. Kennicott (1858) mentioned a Wiscon- 1939, p.131; Osgood, 1936) for the offspring.

396 THE WILD MAMMALS OF WISCONSIN One mother surprised with her young on the Home Range and Density. Males have ground reportedly let them grab her fur and the larger home ranges and may overlap more dragged them into cover (Seton, 1953). The than one female’s home range. Home rang- family of up to five littermates may nurse with es have been estimated as 12 acres (4.8 ha), milk teeth (June 9) even when each male is 3.7 acres (1.5 ha) in winter (Glover, 1943). larger than the mother. Densities have been estimated by number Studies of weasel metabolism have men- of pelts reported (a quantity varying with market tioned that male weasels do not care for the demand) and studies of tracks in snow. The lat- young. Both parents have been found in a ter method depicts home range in winter to some nest, which is as large as a “peck” (Seton, extent. In Wisconsin little has been done, and 1953). The astute mammalogist William one must turn to other states and different races. Hamilton believed that males brought food Polderboer et al. (1941) found one per 40 acre to the young. Some suggest that for a while (16 ha) area, and other studies show higher val- the male brings food to the female. Hall (1951) ues, as much as one per six acres (2.4 ha), and, never observed a male present with young, not surprisingly, some densities much lower. but cited observations by Green that a male Curiously, in the UW-SP museum collec- helped the female remove young from a dis- tion the long-tailed specimens (see below) are turbed nest. Hall believed males may stay with almost all males. In 30 years only a single fe- the mother until the young are partly grown. male was taken, and until it showed up I told Mortality. Raptors such as owls (Errington students in jest the long-tailed weasel is unisex- et al., 1940), large snakes, domestic cats and ual. Some reasons for a great preponderance dogs, foxes, and other carnivores prey on of males that come to mind, determined from long-tailed weasels (Jackson, 1961; Sheffield, other weasel studies, include 1) males wander 1997). Automobiles kill many, and trapping more, in search of females; 2) being larger with is a major cause of mortality. longer bounds males range farther distances, in Jackson (1961) lists a variety of fleas, ticks, search of larger, faster prey such as rabbits; 3) lice, Alaria taxidae, Taenia, Filaroides, Phys- females tend to spend more time in burrows aloptera, Capillaria, and the nasal parasite instead of crossing highways, either in their dens Skrjabingylus nasicola as infesting long-tailed or in burrows of mice; 4) females suffer higher weasels. Sheffield (1997) greatly adds to this natural mortality because of the costs of being list, including also chiggers and mites. mothers or being smaller, and possibly are more easily captured by raptors and carnivores. I have seen female specimens preserved in fair num- Table Car-14Car-14. Cranial measurements of adult long-tailed bers from Beaver Dam and in southeastern lo- weasels. Specimens from Field Mus., Michigan, Michigan calities in the Milwaukee Public Museum. Some State, and UW-SP.* of these were exceptionally small. Locality Total Tail Hind Cond. Cranial Max. Remarks. The long-tailed weasel is a re- l. foot l. br. t-r lentless hunter capable of killing much larger Females animals than itself. It kills mammals such as Upper Mich 335 114 40 43.2 19.3 12.4 the cottontail rabbit, which it pursues by sight Upper Mich 340 114 36 43.5 17.9 12.3 and smell and finally captures with a dash and Oconto Co — — — 40.6 19.0 12.0 tenacious biting of the neck or base of skull. Males Iron Co 435 153 52.5 22.2 15.1 Mice are much more easily dispatched. This Portage Co 420 134 44 48.9 21.2 14.5 weasel will climb trees in search of squirrels, Adams Co** 360 119 43 45.0 21.4 13.0 and doubtless kills some of them. There are * Ratios of tail/head & body > 49, N = 7. at least three vocalizations known (Sheffield, **Age subadult 1997; Svendsen 1976).

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 397 On the Teddy Roosevelt ranch in North 1796. Mustela mink Peale and Palisot de Beauvois. A Dakota, a man kicked at a long-tailed weasel, scientific and descriptive catalogue of Peale’s Mu- which reacted by climbing up his coat and seum, Philadelphia, p. 39, type from Maryland. tenaciously biting him on the throat. After 1854. Putorius nigrescens Audubon and Bachman. The being dislodged, it returned to try it again, viviparous quadrupeds of North America. 3: 104. until the pet bulldog took action (the gaping No type selected; mink were discussed from Penn- men did not) and quickly killed the weasel. sylvania, New York, Vermont, and Canada. Wight (1932) also reports a weasel attacking 1858. Mustela winingus Baird. Mammals. Reports. a man. Another longtail fell to earth together Expl. Surv... 8: 177. with a hawk killed in flight. A poultry killer 1911. Lutreola vison borealis Brass. Aus dem Re- entered a 7/8 inch hole to kill a farmer’s chick- iche der Pelze, p. 504, type from northeastern ens, and left by the same tiny opening. Obvi- North America. ously this was a small female (Seton, 1953), 1999. Neovison vison (Schreber, 1777). If M. lutre- or perhaps subadult. ola signicantly differs from the American minks, Additional natural history. Sheffield and one uses Neovison vison for the American Thomas (1997) studied the long-tailed weasel. mainland species and N. macrodon for the sea Geographic Variation. There is only one mink, or one may assign American minks to geographic race in Wisconsin. the subgenus. Sealfon (in press, J. Mammal.) Specimens examined. Total, 34. Adams, followed Wilson and Reeder’s (2005) summa- Ashland, Clark, Dane, Door: Gill’s Rock 1. ry of chiefly biochemical characters (1: 618-9). Florence, Fond du Lac, Forest, Kenosha, This author agrees. Manitowoc, Marquette, Milwaukee: Milwau- kee 3 Mil. P. Mus. 5 mi. E Brookfield 1. Whitefish Bay 1 MPM. Oconto, Oneida, Neovison vison letifera (Hollister) Pierce, Portage, Waushara, Winnebago counties, Not found: Broggeville 1 MPM. 1913. Mustela vison letifera Hollister. Proc. U.S. Michigan. Alger, Gogebic, Iron, Marquette, Nat. Mus. 44: 475, type from Elk River, Sher- Schoolcraft counties. burne Co., Minnesota.

The name Mustela means “weasel,” and Genus Neovison vison may mean weasel, based on weso, or Baryshnikov & Abramov something similar in German or Scandinavian. The French word for mink is vison today, and Neovison vison (Schreber) the word mink is from the old Swedish word American Mink maenk. Early in the fur business the French in Canada commonly used an Indian word otay. “Messrs Phillips & Woodcock of Cancadea, New This weasel-like, semi-aquatic mammal is larg- York, conceived two years ago the business of er than weasels and it does not molt white. The breeding mink for their fur [on farms, in pens]. Mink subspecific letifera means “bringer of death.” raised for catching rats or for their fur were kept in Description. Resembles a large weasel large pens, and were notably docile and considered with short, round ears, the mink is chocolate pets; they were sociable to one another, except brown overall (giving its pelt an attractive uni- breeding males.” — Elliott Coues, 1877, Fur- color). The throat and chin show splotches bearing mammals. or a patch of white. These vary. The eyes are black, the vibrissae prominent, and the head 1777. Mustela vison Schreber. Die Saugthiere... pl. is held high. If attacked, the screeching, musk 127b. Type from Canada (= Quebec). emissions, feints and overt attacks, and in-

398 THE WILD MAMMALS OF WISCONSIN credible quickness make this little carnivore guard hairs long and oily. There are two intimidating. The long (over a third of the to- molts each year (Rust et al., 1965). The bac- tal length) tail is relatively shorter than that of ulum is sword-shaped (“handle with blade”) the semi-aquatic muskrat. The mink’s tail is having a distal hook and a grooved ventral bushy. When away from water, and mink oc- surface (Elder, 1951; Long and Frank, casionally leave it entirely, the mink feeds on 1968). There is an incredibly variable os cli- birds and mice. When in the woods it resem- toridis in some females (Long and Shirek bles the American marten. The marten has 1970). The feet are semi-webbed (Lowery, shaggy fur, ochraceous (orange) spots on 1974) and fully furred, excepting the black throat and chin, large, erect ears lined on their footpads. On each foot there are five toes, inner surface with buff hairs. To clinch the but the inner one is very small. Chromosomes identification, if a specimen is in hand, the are 2N=30 (Hsu and Benirshcke, 1968). dental formula differs. The mink has 34 teeth; There are anal scent glands. marten and fisher, 38. The difference derives As in most mustelids, the males are about from the number of premolars, 3/3 in Mus- 10-15 percent larger in external measure- tela and Neovison and 4/4 in Martes. The ments, and the weights much greater in males. skull of the mink can be distinguished from Adults vary from 580 to 700 mm (23-28 inch- marten skulls (which are similar in size) by the es) in total length in males, tail 190 to 230, number of premolars, and from skunks (Me- hind foot 68-80, and weights to 1.6 kg (3.6 phitis) by flattened auditory bullae and sub- lbs). Females vary from 460-575, 150 to 190, quadrate upper molar in skunks. The mink, 60 to 70, and have weights up to 1.8 kg (2.4 as in other Mustela and the martens, has lbs). One of the largest specimens of Wiscon- dumbbell upper molars. sin mink, from Ripon, Wisconsin, was a ter- The fur varies from rich reddish brown restrial male with its stomach full of juncoes. to muddy grayish or yellowish brown, and Its external measurements are 656-200-71-25, probably due to the gene influx of released wt. 1.96 kg (4 lbs, 5 oz).Some cranial mea- or escaped ranch mink. There are some surements of 3 adult males and 2 females are, white (with black eyes) and some gray pelag- resepectively, as follows: condylobasal length es. These have trade names in the fur busi- 72.2, 60.6, 66.4, F 56.9, 55.5; interorbital ness, and the colors are inherited. The nor- breadth —, 13.5, 14.0, 12.5, 13.1; zygomat- mal underfur is soft and dense, and the brown ic breadth —, 35.1, 35.1, 32.3, 32.0; cranial breadth —, 30.3, 31.9, 28.8, 26.8; maxillary tooth-row —, 19.8, 21.4, 18.0, 17.7. Dental Formula. DF = I 3/3, C 1/1, P 3/3, M 1/2= 34. Seldom are there dental abnormalities. Geographic Distribution. Usually the mink is found in wetlands or their proximity throughout the state, uncommon in urban areas unless large rivers are present. Some records are of released or escaped mink farm animals, which eventually may change the genetic composition of Wisconsin animals. The mink is more common in the northern counties, where wetlands are more numerous. The animal is widespread in wetlands of Skull of Neovison vison. temperate Canada and the United States.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 399 Status. The mink is an important fur species, even in today’s depressed markets. Usual- ly about 20,000 to 40,000 mink are trapped each season in Wisconsin. Wild mink usually are more valuable than ranch mink, but the fundamental determinant of value is the primeness and luxuriousness of the pelt. Male pelts are more valuable than the smaller pelts of females. Presently the average price of mink among the aquatic furbearers is exceeded only by otters. Protected by state trapping laws, the species is holding its own. It is a predator of ducks and their eggs and muskrats (which are impor-

Maps showing geographic distribution of Neovison vison in Wisconsin and North America.

400 THE WILD MAMMALS OF WISCONSIN tant furbearers), and feeds on fish and frogs. two half-grown mink foraging together, but These predatory actions are therefore injurious usually the mink is solitary. Adult males may to humankind. By their own territorial spacing even fight to the death (Errington 1943; Long and the activities of humans (trapping, habitat and Howard, 1976). Mink are not known to removal), the mink is seldom abundant enough cache foods. to create much of a problem. Although infre- The mink is obviously carnivorous on quently observed the mink is not rare. terrestrial and aquatic prey, and is quite op- There were more than 200 mink farms portunistic from one place to another. Sea- in Wisconsin, the highest number in America sonally, Waller (1962) found predation on (Fruth, 1986). Today there are many animal mammals lowest in spring and highest in sum- rights activists that morally oppose cruelty of mer. Mammals included Peromyscus, Micro- wild animals. In some misguided protests, tus, Ondatra, and Sylvilagus. Birds were tak- people, carried away with zeal, but ignorant en throughout the year. Waterfowl were tak- or without any concern for tragic conse- en, ducklings and adults alike. Crayfish were quences of the released animals, vandalized most abundant in the diet in summer in Iowa. mink farms and have turned out thousands of In Missouri males ate more muskrats, where- domesticated mink into the farmlands (espe- as females preyed more on mice. Errington cially in 1998-2002). This happened in Min- (1943, 1963) determined that problems (hun- nesota, Iowa, and Wisconsin. ger, disease) for muskrats increased the pre- Habitats. The mink dwells usually near dation by mink on them. Other reports on water, regularly ranging away from water as foods are tabulated in Table Car-15. much as 100-200 feet (30.5 to 61 m) to find Reproduction. The mating season is in late food. Occasionally a mink seems to leave the February to early April (Enders, 1952). Matings water entirely. Ordinarily any wetland will do, are promiscuous. During copulation, which is from the bays of Lake Michigan isles, to the famous for its hostility and duration, up to two marshes and swamps of the North Woods, to hours duration, and the male may ejaculate sev- rivers, streams and brooks, and even to shal- eral times. The male seizes the female by the low, marshy drainages. Aquatic foods are the nape and may cause injury. The females remain reasons mink live near water, sometimes nest- in estrus throughout the breeding season, and ing in lodges and burrows of muskrats and bea- ovulation is induced by copulation. Females are vers, and in any available holes and hollows, receptive about every week, and more than one rock and brush piles, log jams, and tree roots. male may father “kits” in a litter (Eagle and Jackson’s (1961:355) and Kennicott’s com- Whitman, 1987). Gestation is about 40 to 70 ments on extensive burrow systems notwith- days, depending when the embryos implant. The standing (see Hoffmeister, 1989) even those average is about 51 days. Thus, there is some mink ranging far from standing water probably delay in implantation, depending on photope- do very little digging. In large burrow systems riod, but not for any long period of time (End- of the muskrat the two species may cohabit the ers and Enders, 1963). After implantation, the burrow (Eagle and Whitman, 1987). Often two young are born about a month later, in late April mink dens are used “simultaneously” [to the or early May, even June (Eagle and Whitman, extent possible]. A mother with “kits,” referring 1987). Litter size varies from 1-8, as many as to young mink, may use the same den up to 40 10 (Jackson, 1961), averaging four. There is days. Subsequently a den may be used for 3-13 only one litter per year. days before departure. The female’s natal nest Newborn mink are nearly naked, blind, is constructed of grasses and feathers. and helpless at birth. There is a fine coat of Foods. Mink usually forage at night, but whitish hairs dorsally. Each neonate weighs are seen hunting in the daytime. I have seen 8-10 g (0.28-0.35 oz). Deciduous teeth erupt

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 401 between 14 and 21 days. The eyes open at These PCB’s were obtained from purchased three weeks. The young then begin to eat solid fishes used as mink food. Mercuralism in mink food. By seven weeks of age they are 60 per- is an important source of mink mortality (Lin- cent grown, and play with one another, scombe et al. 1982; Eagle and Whitman 1987). squealing, and attacking, until weaning takes Home Range and Density. A study on place in late summer (Linscombe et al., 1982). home range was made in Sweden (Gerell, Only the mothers care for the young. After 1970) by use of telemetry. He found range 5-6 weeks the kits are weaned. Mink mature for adult males to average 2,630 m (1.63 mi) at approximately 10 months of age. Females in stream length, whereas juvenile males var- attain adult weight their first autumn, males ied from 1,050 to 1,400 m (0.62-0.87 mi). in their second year. Longevity is 5 years, Adult females averaged 1,850 m (1.15 mi). reportedly as long as 10 (Crandall, 1964). Males may have a home range twice as large Mortality. Trapping is a major cause of as the home range of females. Home range mortality of mink in America today (and has is restricted in winter. When young of the year been since the 1920’s), even though markets dispersed, some movements were extensive. for fur have fallen. Russia has become the One male moved 21 km (13 mi) in 27 days; top mink fur producer in the world. Wiscon- another moved 45 km (28 mi). The female sin is one of the top mink fur states. Protec- may linger near her den. tion of wetlands should help this species in Shoreline males occupied 2.5-5.5 km Wisconsin. Banfield (1974) found trapping (1.5-3.5 miles), and females 0.5-3 (0.3-1.9 drastically reduces abundance of mink. Natu- miles). In marshes home ranges of males were ral predation is probably not so important as 316-1626 ha (781-4018 acres). Home rang- disease, hunger, and other causes of mortal- es are small in winter. During courtship, home ity, especially trapping. Occasional predators ranges in males are 16-34 % larger (Eagle include bear, fisher, red fox, gray fox, bob- and Whitman, 1987). cat, lynx, wolf, and great horned owl (Lin- Densities are usually determined by the scombe et al., 1982; Eagle and Whitman, fur catch, which usually mentions only the 1987). Mink are occasionally road-killed. numbers per county. Such estimates also suf- Internal parasites include tapeworms, fer because of the demand (prices) for mink flukes, many roundworms, the spiny-headed pelts, and inexplicable rise and fall of the num- worm, and sporozoans (Isospora) (Linscombe ber of trappers regardless of price. Densities et al., 1982). They also list external mites, fleas, seem highest in cattail marshes where there and ticks. A student showed me a bear tick, are many muskrats (Errington, 1943). Err- the carrier of Lyme disease, attached to his ington found 9-15 per square mile. Marshall thumb, contracted from a snap-trapped mink. (1936) found 1.5 per square mile in Michi- Dr. Stephen Taft showed me many large kid- gan. Linduska (1950) found populations as ney worms Dioctophyma, almost always from high as 4 per square mile the mink’s right kidney, that his student Con- In Wisconsin, McCabe (1949) found 24 nie Frederick collected in wild Wisconsin mink. mink present in a marsh with a density of 1/ Various diseases affecting mink populations 18.8 ha (1.25/square mi). Subsequently the include botulism, anthrax, tularemia, tuberculo- number of mink declined to 7-10 annually. sis, distemper, and many others, none of which Remarks. The mink is a swift swimmer has been noted in wild mink populations. often capable of overtaking fishes. On the land Pollutants cause harm to wild mink. Both it lopes along with its body humped high in ranch and wild mink are affected in their repro- the middle, or dashes weasel-like in sudden duction (of “minimink”, dwarfed young often runs. It tends to wrap its long body about its aborted) by polychlorinated biphenyls (PCB’s). prey as it administers the fatal bite.

402 THE WILD MAMMALS OF WISCONSIN Mink ranches. In Wisconsin there are and physiology of mink has been determined many mink ranches, although the number has from these animals often inbred, fed on the fallen in recent years because fur markets were same diet, killed at one time, with age and depressed over the past few years. There are sex closely controlled. signs of renewed demand for fur. Other coun- Additional natural history. Lariviare tries compete with the United States in mink (1999) reviewed biology of the mink. production, including Russia. Some large Geographic variation. There is a single Wisconsin minkeries produce 40,000 pelts subspecies in Wisconsin. per year. Much information of the anatomy Specimens examined. Total, 33. Craw- ford, Dodge, Fond du Lac, Green Lake, Iron, Jefferson, La Crosse, Manitowoc, Marinette, Table Car-15. Foods (percentages) of Mink in Winter in Oconto, Oneida, Outagamie, Portage, Rock, Missouri and Michigan. Sheboygan, Trempealeau, Walworth, Korschgen Sealander Waukesha, Winnebago, Wood counties. Missouri Michigan Other records (After Balliett and Taft, Food Occurrence Vol. Vol. Occurrence 1978): Chippewa, Monroe, Waupaca, Price, Mice& rats 22.6 23.9 15 11 Fond du Lac, Marathon counties (Not plotted). Rabbits 5.9 10.2 15 14 Muskrats 1.1 1.3 36 31 Frogs 25.5 24.9 23 20 Genus Mephitis E. Geoffroy Saint- Fishes 30.9 19.9 11 6 Crayfish 19.9 9.3 6 2 Hilaire and Cuvier Birds 5.9 5.6 13 15 Striped and Hooded Skunks Inverts 4.0 0.4 — — Snakes 0.3 0.1 2 2 Known formerly as Chincha Lesson, 1842, the older name Mephitis, named in 1795, has priority. The Mephitis are medium-sized carnivores, larger than the eastern spotted skunk, Spilogale, and stout of form. Striped skunks have flattened auditory bullae and a shallow step on the inferior surface of the dentary. The face is downturned from the cranium, so that in profile there is a fairly sharp angle above and just behind the orbits. In Spilogale the rostrum and the cranium form nearly a straight line. Skunk fur is black and white, an advertisement or warning coloration. The scent glands are famous for their powerful odor. The palate terminates just behind the upper molars; the posterior molar is both elongate and broad (subquadrate) and the upper carnassial has a posterior shearing edge confluent with a sharp, piercing anterior cusp. The skunks show resemblance to otters, bad- Sketch of upper carnassial and upper molar of the mink gers (especially stink badgers Mydaus), and in Neovison vison and skunk Mephitis. The difference between some features (e.g., baculum) seem divergent musteline and mephitine teeth is egregious, but compare the from the other mustelids. Dragoo and Honey- latter with otter’s and badger’s skull photos. The bony ear of otter and skunk is also similar. These characters help in cutt (1997) proposed to raise the skunks and identification and should in cladistics. Long (1976c, 1981). stink badgers to family status, Mephitidae. They

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 403 might have ignored some of the morphological or its skin may be identified by its black and evidence showing stink badgers are badgers, white color pattern, beginning as a narrow, and that skunks resemble otters. median pin-stripe of white on the head, and expanding as a broad band of white behind the ears along the nape of the neck. This band Mephitis mephitis (Schreber) divides as two nearly parallel stripes of white Striped Skunk posteriorly. They terminate on the back or rump, but sometimes extend into the bushy “Strong ammonia, essence of garlic, burning tail fur several inches. White hairs may or may sulphur, a volume of sewer gas, a vitriol spray, a not be intermixed in the black tail hairs. The dash of perfume musk, all mixed together and remainder of the animal is black or dark black- intensified a thousand times.” — Ernest Thompson ish brown. This description will not work if Seton, Lives of Game Animals. the skunk is all black, or if the stripe pattern is abnormal. 1776. Viverra mephitis Schreber. Die Saugthiere... The form of the body is stout, rotund, Theil 3, Heft 17, pl. 121, type from eastern appearing plump (and indeed the skin is usu- Canada (= Quebec). ally layered with fat) with fairly long hind legs 1902. Mephitis mephitis: J. A. Allen et al., Science, (elevating the hinder end when the skunk n.s. 16: 115. ambles along, and elevating the anus when 1818. Mephitis americana Desmarest. Nouv. Dict. the skunk threatens or emits scent). The fore- Hist. Nat. Paris, 21: 514, no type. Possibly paws have elongate curved black claws. Ob- this name applies to this species. viously a digger, the toes are slightly webbed as in badgers. The feet are pentadactyl and The word skunk is of Native American plantigrade. The eyes are black. There is no derivation. The word is derived from the Ab- known nictitating membrane, as seen in the naki segak. Latin Mephitis means having a badger Taxidea. foul odor. The odor is emitted by a pair of anal The skull is about the size of that of mar- scent glands encapsulated by strong muscles. ten or mink; they are described in their ac- A yellowish liquid emission with powerful odor counts. Briefly: the striped skunk has flattened can be directed at an adversary accurately, even auditory bullae, the shape of the carnassial is occasionally at 6 m. This odor is primarily for pointed, with a long posterior shearing edge, defense, and the liquid can temporarily cause the subquadrate upper molar is robust, and blindness. Only stink badgers (Mydaus) rival there is a short hard palate. The skull resem- skunks for emitting such foul odor. bles that of Spilogale but is significantly larger (see account of Spilogale). There are six or seven pairs of mammae, Mephitis mephitis hudsonica Richardson occasionally more (range 10-15). The bacu- Striped Skunk lum is a peculiar slender spicule (Burt, 1960). Chromosomes have a diploid number of 50, 1829. Mephitis americana hudsonica Richardson. FN = 92 (Wade-Smith and Verts, 1982). Fauna Boreali-Americana. 1: 55, type from The color pattern is advertisement, for plains of the Saskatchewan, Canada. intimidation and warning of the foul scent 1911. Mephitis minnesotae Brass. Aus dem Reiche der emitted (which also burns the eyes). The bold Pelze, p. 532, type from northern Minnesota. pattern is seen in newborn skunks even before hair grows out. The underparts are en- Description. See remarks above in the tirely black or brownish black, with an occa- account of Mephitis. The animal in the wild sional white spot on the venter. There is an

404 THE WILD MAMMALS OF WISCONSIN annual molt, beginning in April with (1983) legions in the southern and western prairie coun- lists total length as 520-680, tail length 180- ties. They are found in fairly large cities, and 280, hind foot 64-80, and ear length 25-35 may enter any of the villages or small towns. for males, and for females, 520-650; 175- They seldom come into contact with people, 270; 57-76; 25-35. He reported fat males in but they may encounter the pet dog. See Map. autumn weighing up to 12 pounds (5.4 kg). Status. Judging by road-kills the striped Although females are reportedly smaller than skunk and the raccoon (Procyon) are the most males, Verts (1967) found a great deal of over- abundant carnivores in Wisconsin today (also lap in his samples of the two sexes. From Table Car-1). The striped skunk has relatively Wisconsin, adult males (2) and females (2) had little value as a fur resource, because the un- measurements as follows (ear length omitted): popular connotation of and association with total length 545, 615, 525, 510; tail length skunk fur depresses the value. Skinning a skunk 50, 70, 63, 65; hind foot 195, 185, 225, is not pleasant. In spite of carcasses wasted, 202; condylobasal length —, 75.5, 70.9, the furs harvested in Wisconsin provide mon- 68.5; zygomatic breadth —, 46.1, 43.0, —; ey in the hundreds of thousands of dollars when interorbital breadth —, 21.7, 20.5, 20.0; all the numbers are counted. Rolley (1996) maxillary toothrow —, 23.0, 22.4, 20.3. reports that the number of furs taken annually Dental formula. I 3/3, C 1/1, P 3/3, M is stable, and has been since 1986. 1/2 = 34. The greatest value of the skunk to human- Geographic Distribution. The striped kind is as a predator of small mammals and skunk occurs throughout the state, except on other injurious pests. These things have been some islands in Lake Superior and Lake Mich- said against skunks: they are a reservoir for igan. It occurs on Washington Island, report- rabies virus, kill chickens, eat bees, dig holes edly released by a fur breeder years ago (Long, on lawns and golf courses, kill ground-dwell- 1978). Jackson (1961) without comment ing birds, and eat their eggs. To these nega- mentions a record from the Apostle Islands. tives can be added the foul odor, and the emis- This skunk also occurs widespread in Upper sion of it onto an unwary person, or the pet Michigan (Baker, 1983). dog. In their defense, Jackson mentions that One may see road-killed skunks on the they seldom kill chickens, and never those in highways from Maiden Rock and Ellsworth in well-built pens. They seldom eat bees, never if the west eastward to the Door Peninsula, and the hive is set several feet above ground. They even on Washington Island. They are killed in do dig holes but do so in search of cutworms and other pests, doing a favor in the long run. They seldom eat birds. Many chickens and birds eaten are carrion. A skunk that becomes a persistent pest may be trapped with a humane trap and removed. Jackson (1961) writes, “One should never wantonly kill skunks.” Rabies is at this time a real problem. The poor skunks presently suffer epidemics of rabies, being afflicted more than any other Wis- consin mammal (now that dogs are innoculat- ed with vaccine). About half of the animals that tested positive for rabies in the State Hygiene Laboratory (see Figure) are skunks, and the true incidence in wild skunks may be even high- Skull of Mephitis mephitis. er. Skunks seldom bite people unless handled.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 405 If a skunk bites someone, the rabies vaccine must be administered. Expect in the future broadcasting baits with oral vaccine to reduce the incidence of rabies in skunks. Habitat. The striped skunks inhabit many kinds of habitats, wet to dry, rocky to prairie soils, coniferous forest to hardwoods, woodlots, weedy prairies, brushy areas, hedgerows, grassy roadsides, wetlands of all kinds, ravines, gullies, and almost any place where cover is sufficient and mice and insects available for food. Young of striped skunks are reared in natal dens (Verts, 1967) and often use other

Maps showing geographic distribution of Mephitis mephitis in Wisconsin and North America.

406 THE WILD MAMMALS OF WISCONSIN Sketch of striped skunk feeding on turtle eggs. Lloyd Graph showing rabies incidence in Wisconsin and the Sanford. United States in skunks. mammals’ burrows (73 % of them that were rial in 92 %, and other animal matter in 95 dug by badgers, 17 % by red foxes, 7 % by %. The diet, then, is not strictly insectivorous the skunks themselves, and 3 % by muskrats). and carnivorous, but in season the skunk eats They also den in woodchuck dens and rabbit corn, cherries, nightshade, and other fruits. burrows; in Wisconsin such use would seem Worms are difficult to detect, but doubtless likely. Most of the growing season, skunks eaten, as are crayfish, turtle eggs, eggs and live in temporary outliers above grounds, and young of ground-nesting birds, and reptiles in winter a more secure place to sleep is used, and amphibians. As mentioned above skunks such as a hollow log or stump, or underground eat many small rodents. burrow (Crabb, 1948). In winter the body tem- In Michigan, Dearborn (1932) found in perature falls in northern, cold latitudes, but digestive tracts and fecal scats (mostly collect- skunks do not hibernate. In southern Wiscon- ed in summer) insects 57.3 % frequency; fruit, sin they often wander about in winter, but in 17.7%; grain 12.1%; small mammals 10.2%; Upper Michigan they seldom are out on the birds, 2.35%; turtle eggs 0.21%; bird eggs snow (Baker, 1983). 0.11%; reptiles 0.01%; and nuts 0.01%. The Up to three nests have been observed with- small mammals eaten were local kinds, Per- in the burrow (Allen and Shapton, 1942), and omyscus, Microtus, Zapus, house mice and there may be as many as five entrances. Nests Norway rats, several kinds of squirrels, moles were large and bulky, consisting of dry grasses and shrews, cottontails, raccoons, and wood- and leaves. Six nests were found above ground chucks (probably the larger mammals were in hayfields. The mother may move old bed- young or carrion). Fishes, frogs, lizards, and ding out prior to making a natal nest. In winter, snakes were also eaten. Adult males eat young two or three (of both sexes) may den up togeth- skunks. In winter obviously the fruit declines er, and ten have been found in a den (Baker, and the numbers of mice rise. In summer, 1983). Study on striped skunk dens in Michi- many insects are eaten, mostly those that live gan, especially winter dens, was accomplished in the soil or on grasses (Hemiptera, Co- by Allen (1939) and Allen and Shapton (1942). leoptera, larval cutworms, crickets, and grass- Natal and winter dens are often excavated or hoppers). Clams, crayfish, and earthworms appropriated, after abandoned by other mam- are opportunistically eaten, and garbage is mals, and farm buildings are often utilized. often eaten. On the negative side, corn, poul- Foods. The striped skunk is an omnivore. try and eggs, garden berries, and honey are Verts (1967) analyzed scats in Illinois, finding taken in raids on the farmyard (Storer and insects present in 88 % of them, plant mate- Vansell, 1935; Schofield, 1957).

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 407 Reproduction. Mating occurs normally in include gray wolf (= timber wolf), coyote, fox- February and March (Verts, 1967; Allen, es, fisher, bobcat, and lynx (Baker, 1983). 1939). There is induced ovulation, about 42 Parasites have been listed by Lawrence hours after copulation. The gestation period et al. (1965) and Verts (1967). They include seems variable, ranging from 59 to 77 days. a flea, a chewing louse, a tick, and a mite in Therefore, there seems some variable delay Michigan, and in Illinois other fleas, flukes, in implantation, as in other mustelids. Young tapeworms, 19 kinds of roundworms, four are born in May or early June. Litter size in kinds of spiny-headed worms, and even a northwestern Illinois was 6.3 offspring (range pentastomid. The nasal passages are infested 1-9). Baker (1983) reports a maximum of 10 with the roundworm Skrjabingylus as are embryos; most females resorbed some em- weasels and the mink. Baker (1983) mentions bryos. There are reports that a few skunks a cuterebrid (botfly larva). Diseases include may mate again in May and produce a sec- pneumonia, tularemia, leptospirosis, histo- ond litter (Seton, 1953; Verts, 1967). Usual- plasmosis, canine distemper, and, of course, ly the late conception was from a mother that rabies (Verts, 1967). earlier did not conceive, lost her litter, or did Home range and density. Males have larg- not lactate (Wade-Smith and Verts, 1982). er home ranges than females, and both sexes Young-of-the-year do not breed. wander about more in late summer and fall The young weigh about 28-35 g, and are (Verts, 1967). In summer wanderings, the born with the eyes closed, the ears closed, skunk seldom exceeds a mile in foraging, and and with fine hairs on a boldly pigmented skin. probably moves about even less in winter, ex- In three weeks the young weigh about 110 g cept on warm nights. In the breeding season (3.85 oz). The ears unfold in 8 days, and the males may travel 4-5 miles (6.4-8 km) per night eyes open in 22. By then the young are furred (Edmunds, 1974). In mixed farmland there may out. The young weigh about 220 g (7.7 oz) be only one per 100 acres (40.5 ha) or as many by six weeks. About then the young eat solid as one per 10.4 acres (4.2 ha). food. Weaning occurs at 46-49 days or soon- Remarks. Skunks have been raised on er. The young wander about with the mother fur farms and kept as pets (Seton, 1953; Jack- much like Meles, with outliers often used for son, 1961). Young may have the scent glands temporary quarters. Some lucky observers are surgically removed (Bailey, 1937). have seen the mother leading her young sin- To decrease the incidence of rabies that gle file. By August the young leave the moth- has developed in wild skunks, they have been er. Animals attain adult size by 10 months, gassed and killed locally in large numbers. and may live 6-7 years. These practices are costly, rarely effective, and Mortality. Humans are the chief enemy, sometimes inhumane. not only by trapping hundreds taken each year, Striped skunks are not aggressive (unless shooting (with no way to estimate how many rabid) and to provoke the defensive posture are illegally shot, as skunks are often misun- and/or emission of scent usually the skunk derstood and detested), and others killed by must be attacked, surprised, or cornered, as thousands on the roads. Insecticides and her- one was in my garage by the dog. On the bicides may also take a toll. Large owls will eat highways, unfortunately for the skunk and the them, and some large carnivores will likewise. driver, the fearful skunk will cease fleeing, turn Chuck Pils (personal correspondence) observed to threaten the oncoming car with its defen- a striped skunk, still writhing, carried over a sive posture (presenting the anal glands), and road in Iowa County by a great-horned owl. many are killed in this manner. The skunks Badgers may eat striped skunks on occasion are usually solitary except in the mating sea- (Long and Killingley, 1983). Other predators son, and are nocturnal, occasionally abroad

408 THE WILD MAMMALS OF WISCONSIN in daylight. The activities are centered about nigra. Verts (1967) found remarkable varia- the den site (Allen, 1938). tion in the incidence or frequency of patterns Additional Natural History. Verts, 1967, in northern Illinois, which may be where in- and Wade-Smith and Verts, 1982, reviewed tergradation is occurring between north and the biology of the striped skunk. south. I follow Jackson’s (1961: 376) opinion: Geographic variation. The taxonomy of “There is great variation in characters... but the geographic races is badly in need of a com- this variation cannot be assigned either to any prehensive revision. One must start with A. particular geographic range or any other group- H. Howell’s monograph in 1901. Howell ing. About 10 per cent of the specimens from worked in a time of little regard by taxono- southeastern Wisconsin (Racine, Walworth, mists for narrow or broad zones of intergrada- and Rock counties) show some trend in char- tion. In those days even the intergrades were acters toward M. m. avia, but on the whole often given subspecific names. Hall and Kelson are nearer to hudsonica.” I assign all Wiscon- (1959) arbitarily mapped a boundary between sin specimens of striped skunks to M. m. hud- two supposed races along (but not identical to) sonica, for there is no evidence that those in the Illinois border, with some Wisconsin records the south differ appreciably from those in the assigned to a southern (avia) and others to the north. There is evidence of intergradation in northern hudsonica. Neither determination northern Illinois, where a true boundary prob- was based on examined specimens. Jackson ably will depict hudsonica entering Illinois. (1961), with some deference to Howell’s work, Specimens examined. Total, 22. Ash- assigned all Wisconsin skunks to the northern land, Dane, Door Co.: Washington Island, race. Hoffmeister (1989) commented that un- Deer Swamp (no specimen). Dodge, Dunn, til a modern, objective taxonomic revision was Juneau, La Crosse, Lincoln, Manitowoc, made he would assign all the Illinois specimens Marathon, Oconto, Portage, Rusk, Sha- to a southern race. Thus, we see here another wano, Taylor, Waukesha, Wood counties. example created of what I have termed Jack- son-Hoffmeister phenomena, where the political boundary between Illinois and Wisconsin Genus Spilogale Gray delineates the boundary between geographic Spotted Skunks races. Baker (1983) assigned specimens from lower Michigan to eastern M. m. nigra. Small, black-and-white skunks with usually If the race M. m. mephitis from eastern four, and as many as six white stripes, some Canada significantly differs from the Maryland of which may be broken up as series of spots race M. m. nigra, I would not be surprised. If on the upper parts. Tail length less than length there is a wide-ranging race along the Rockies of head and body. The body is slender, not and extending far northward, I would not be stout as in Mephitis. Hard palate of mouth surprised. Whether there exists in Illinois a race extending posteriorly slightly beyond the up- ranging northward to and possibly beyond the per molars. The inferior margin of the den- Wisconsin boundary, and ranging westward to tary is slightly curved but lacks the step seen Kansas, as a differentiated prairie subspecies, in Mephitis. The color is warning coloration, needs verification. The Mississippi River and advertising the ability to throw a liquid and the Great Lakes (see Baker, 1983, for further gaseous emission with a powerful, acrid and discussion) might have provided sufficient geo- disgusting odor upon its adversary. This may graphic isolation for speciation. be facilitated by the “hand stand”, when the M. m. avia is of dubious validity. Hoffmeis- little skunk stands on its forepaws facing its ter (1989) reports the Illinois subspecies aver- adversary from between them, with its anal ages larger size than those typical of M. m. glands and tail directed from a higher plane,

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 409 from an intimidating stance, and possibly with Spilogale putorius (Linnaeus) enhanced aim (C. E. Johnson, 1921). Eastern Spotted Skunk There is a single kind of spotted skunk in the Upper Midwest (Van Gelder, 1959), 1758. [Viverra] putorius Linnaeus. Syst. nat., ed. 10, known as Spilogale putorius interrupta. This 1: 44, type from South Carolina. animal has crossed the Mississippi or St.Croix 1875. Spilogale putorius: Coues. Bull. U.S. Geol. rivers into Wisconsin from either Minnesota and Geog. Surv. Terr., ser. 2, 1: 12. or Iowa, but apparently has been eradicated 1890. Spilogale ringens Merriam. N. Amer. Fauna, in Wisconsin, is in peril perhaps in Minneso- 4: 9, type from Greensborough, Alabama. ta, and declining in Iowa and other states (Bop- 1820. Mephitis interrupta Rafinesque. Ann. Nat., ple and Long, 1994). When inquiring of peo- 1: 3, type from Upper Missouri River. ple, ask about “civets”, for spotted skunks are 1859. Mephitis quaterlinearis Winans. Kansan often mistakenly known as civets. newspaper account mentioned by Coues: Fur- Long (1965) and Mead (1968) called at- bearing animals... 1877. No type designated. tention to the reproductive isolation of the western S. gracilis from eastern S. putorius. Be that as it may, Van Gelder (1959) consid- Spilogale putorius interrupta (Rafinesque) ered the two as one species because of possible intergradation perceived in some speci- 1820. Mephitis interrupta Rafinesque. Loc. cit. mens from Mexico. I mapped the entire range 1952. Spilogale putorius interrupta: McCarley. for the two as one species, but certainly in Texas J. Sci., 4: 108. the United States the two act as good species and have been described separately (e.g, in The name Spilogale means “spotted wea- Wilson and Ruff, 1999). sel,” and putorius means foul smelling. The subspecific name interrupta refers to the tendency of some stripes to break up into spots. Description. The eastern spotted skunk is a small, boldly marked, and beautiful black and white mammal, with short legs, short ears, and fluffy tail. The skull resembles that of Mephitis, with flattened auditory bullae and trenchant upper carnassial, but there is no step on the inferior margin of the mandible. The external and cranial measurments are smaller than in Mephitis. See genus Spilogale above. The foreclaws are sharp and stout. The baculum is a small, curved spicule (Burt 1960). The eastern spotted skunk has white stripes and spots on a black background, with some whitish hairs in the pencil of the tail. Wisconsin specimens are much darker than those of western skunks. A white spot is found on the forehead. Two more white spots are found just anterior to the ears. Spots are found on either side of the base of the tail, and two Skin, dorsal and ventral aspects of skull of Spilogale more set apart on either side of the rump. putorius. Skin courtesy Frank Iwen. The two pairs of longitudinal stripes curve

410 THE WILD MAMMALS OF WISCONSIN downward over the hind quarters, and spots averaging about 1 pound 1 oz. (0.5 kg) may be more or less present in the lateral throughout. The largest female weighed 1 lb. stripes. Some specimens are nearly black. 6 oz (0.621 kg). Males varied from 1 pound There is a single summer molt. There are ten 7 oz (0.65 kg) in summer, to 1 lb. 8 oz in fall, mammae arranged in two parallel rows. to 1 lb. 9 oz in winter and 1 lb. 10 oz (0.75 From Jackson (1961) the measurements kg) in spring. This is the time of courtship. are given as total length 470-550 mm (18.3- Maximum weights for the males were 2 lbs 8 21.5 in), tail length 178-220 (6.9-8.6 in), hind oz (1.232 kg) in February and 2 lbs. 12 oz foot length 48-52 (1.9-2 in), weight up to 1.3 (1.344 kg) in March. kg in males, and in females 445-482, 165- Dental formula. DF = I 3/3, C 1/1, P 200, 43-47, up to 0.6 kg (2.6 lbs) (based mostly 3/3, M 1/2 = 34. on Iowa specimens). The skull seldom exceeds Geographic Distribution. The eastern 50 mm (2 in) in greatest length (range 45-54). spotted skunk formerly occurred in western Thus, males average a little larger than females and southwestern Wisconsin, but is known (about 5 per cent in total length, a quarter only from two localities (see specimens ex- heavier). Iwen’s measurements of his specimen amined below). The Jefferson County speci- from Jefferson County are total length 620, men is surprisingly far eastward from the nor- tail 290, hind foot 50, ear 30 mm, with a mal range of Spilogale, and the species ap- weight of 872 g (30.5 oz). The skull measured parently must have crossed several great riv- 58 mm condylobasal length, zygomatic breadth ers. One would suspect it came from Illinois 37.8, mastoid breadth, 31.9 mm. This speci- prairies, but Spilogale is unknown in Illinois. men is a very large female, and possibly the Apparently the St. Croix specimen made it measured tail included the hairy pencil. across the St. Croix River, or worked its way Some measurements given by Crabb northward from farther south. The Jefferson (1944) show variation of weights with sea- County skunks may have come from the Iowa son, the males being much larger in winter, populations, crossing the Mississippi south of and the largest female was taken in Septem- the Wisconsin River and working their way, ber. The females varied little during the year, perhaps in several skunk generations, eastward. It seems unlikely two spotted skunks would have been released in Jefferson County. To assume the spotted skunks crossed the Mississippi from the west instead of arriving from Illinois, perhaps long ago (since Parma- lee and Hoffmeister reported them from archaeological digs in Illinois, along the Misissip- pi River, see Hoffmeister, 1989) is logical. Not only are the little skunks absent in Illinois in historic times, but their occurrence in Wiscon- sin is coincidental with the northward expansion of this mammal’s range into southern Min- nesota. Land use until recently encouraged the presence of this little skunk, near small farms, woodlots, small sheds, and haystacks. Status. On the west bank of the Missis- sippi River, and westward, the little spotted Maps showing geographic distribution of Spilogale skunks thrived and moved northward proba- putorius in North America. See text. bly owing to favorable habitats caused by land

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 411 use. Not only was the farming then beneficial it feeds extensively on Mus and Rattus. Its fur to the skunks, providing sheds, outbuildings, occasionally has value. It is one of the most fences and fencerows, hedges, wood lots, and beautiful mammals that ever graced the coun- old fields with saplings and shrubs, but also tryside of Wisconsin, attractive both in pattern available prey, an abundance of house mice and luxurious fur. As a boy in eastern Kansas, and Norway rats. In the 1940’s, the Spilogale I trapped one, which my neighbors told me established themselves in northern Iowa was a civet cat (which it surely was not), and I (Bowles, 1975) and invaded Minnesota, reach- remember like yesterday its little body lovely ing a peak (judging by fur reports) about 1945 and soft with rich black and pure white colors. (Wires and Baker, 1994). About this time the The spotted skunk seldom harbors rabies, in spotted skunk crossed the great rivers, either comparison to Mephitis mephitis. the Mississippi near Lake Pepin or below, or Another thing to consider is the nation- the wide St. Croix near Hudson, establishing al problem of preserving rare species. If the a “beachhead” in the prairie hills and goat spotted skunk seems on the decline through- prairies, and the farms and fields of south- out the Upper Midwest, and it does, then why western Wisconsin. N. R. Barger (1951) and not preserve the species wherever possible? Walter E. Scott (1951) reported the first record Minnesota is looking at the problem, but may for the state, from Bass Lake, Sect. 25, T 30 have some trouble re-establishing the little N, R 19 W in St. Croix County. The date was spotted skunk in highly urban Edina. So why Nov. 19, 1946. Frank A. Iwen of the UW- not save a population across the Mississippi Madison University Vertebrate Museum rein Wisconsin? The eastern spotted skunk ported the second and last record of occur- might be reintroduced in brushy prairie pre- rence. This specimen was taken in Novem- serves, with wild bison, Franklin’s ground ber 1955, near Fort Atkinson, Jefferson squirrels, and other prairie and mixed-broad- County, and another skunk was at the den, leaf community mammals. but escaped from the trap of Robert Ward. Habitat. This skunk is a prairie mammal, No other spotted skunk has been found in of the Great Plains, but it is not a pure grass- Wisconsin (Bopple and Long, 1994). land form. It is more likely taken in gullies and The species seems in decline everywhere, along stream banks. The habitats are shrubby especially in neighboring Minnesota and Iowa hedges and fields, grain fields, weed patches, (Bopple and Long, 1994; Wires and Baker, old fields, outbuildings around farms, wood- 1994). A new and different kind of land use lots, and not-too-well kept farms. The spotted seems to be adverse to these mammals. Re- skunk has never been found anywhere in the moval of wild shrubs, plowing and mowing of state except in southwest and western Wiscon- weedy fields and roadsides, modern rat proof sin, with its mixture of riparian lowland decid- buildings, loss of woodlots, and human caused uous woodlands and grassy hillsides or valleys. mortality all decrease the numbers of skunks. The dens in Iowa, source of our skunk Perhaps the loss of a mammal with the population, were described by Crabb (1948) odious persona of skunk is not mourned. Per- in many kinds of dry shelters, such as rocky haps a mammal popularly, although [incorrect- crevices, hollow trees, log or stumps, under a ly] called the “hydrophobia skunk” is one kind stump or windfall, or in a shallow burrow or Wisconsin can afford to lose. Since we had even the burrow of another mammal. only a few, why concern ourselves with its erad- The Wisconsin den was a burrow under ication? To begin with, the spotted skunk is a milk house (Iwen, 1958). Man-made abodes on the whole a beneficial mammal, feeding on are woodpiles, old buildings, barns and gra- mice, small rabbits, and insects the way it does naries, haystacks, and junkpiles. Dens are used (Bopple and Long, 1994). Around barnyards by several skunks sometimes, except the na-

412 THE WILD MAMMALS OF WISCONSIN tal nest which is maintained by the mother. 156 g (5.4-5.5 oz). Playing began at 37 days, An internal, protected nest of grasses, leaves, when the young wandered about like kittens. or other vegetation is where she rears her By 48 days the young ate meat with the moth- young. Temporary dens have no nests. er. By 54 days, the young appeared to be half Foods. Spotted skunks are somewhat grown and weaned. omnivorous, eating fruits in season and some- Mortality. Humankind is the chief ene- times corn and other grains. A study of scats my, by trapping, road-kills, and modern land in Iowa (Crabb, 1941) found mice, cotton- use practices, such as mowing and eliminat- tails, and corn as winter foods. In spring about ing fencerows and haystacks. Domestic dogs 80 percent was Microtis sp., and nearly half kill many, even though this skunk can climb the scats contained insects. In summer 92 some trees. No doubt owls and badgers occa- percent frequency of the scats contained in- sionally eat them. Occasionally the skunk suf- sects, and mammals dropped to only 35 per- fers from rabies or tularemia. Jackson (1961) cent. Fruits, birds, and bird eggs were com- lists chewing lice, fleas, and a tick Ixodes coo- mon in summer. In autunm, insects were pre- ki. Internal parasites include the fluke Alaria, dominant (80.5 % of the scats), mammals up two tapeworms, and four species of round to 58 %, fruits up to 36 %, and birds declined worms (Erickson, 1946). to only 3.78 %. Insects eaten were mostly Home Range and Density. There is little carabids and scarabids. See Table Car-16. known of this little spotted skunk either in Norway rats were common prey, making up Wisconsin or nearby states. The home range for the rare chicken taken by Spilogale. Some expanded in males in the spring of the year. carrion was eaten. Spotted skunks do not Crabb (1948) found densities as high as five cache food. They often return to food. skunks per km2 (13 to the square mile). Reproduction. Rodney Mead (1968) has Remarks. An opportunistic forager (see worked out the several reproductive patterns Foods above), the spotted skunk is not strict- in spotted skunks. The eastern spotted skunk ly solitary and often is found with one or more breeds in the spring (April). It has no apprecia- other spotted skunks. The animal, not active ble delay in implantation, and the young are in winter, does not hibernate. It is active on born possibly in April or May into June. The warm days, when it feeds mostly on mice if litter size is four or five (range 2-7). The young they are available. are born with sparse yellowish fur, eyes closed. Additional Natural History. Kinlaw Crabb (1944) measured and weighed young (1995) reviewed the biology of the eastern born from a wild-caught pregnant skunk. The spotted skunk. parturition date was June 7, and the weight Geographic variation. There is no speci- was 9.5 g (0.33 oz). Squeaking was heard ation evident in this recent arrival. emitted by the young. Sparse hairs covered a Specimens examined. Total, 2. Jeffer- mostly naked body, which showed the pigment son Co.: Fort Atkinson 1 UW (1955). St. patterns of the adult. Eyes and ears were closed. Croix Co.: Bass Lake 1 UW (1948). At seven days, the weights were about 22.5 g (0.79 oz), and the hair was only 2 mm long. Table Car-16. Foods of Spilogale putorius. From scats. At 24 days the young threatened by elevating After Crabb, 1941; see Bopple and Long, 1994. the tail. By 28 days, the weights varied from 116 to 127 g (4-4.9 oz), and the bodies were Foods Winter Spring Summer Fall sleek and fat. At 32 days, the eyes opened, Mammals 90.4% 66.6 35 58.3 some teeth were visible in the gums and weights Birds 5.3 5.8 11 3.7 were 128 to 141 g (4.5-4.9 oz). By 36 days Plant material 31.9 10.6 20.7 56.2 canines had erupted, and weights were 153- Arthropods 14.6 47.6 92.35 80.5

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 413 Genus Lontra Gray The dental formula numbers 36 teeth, differ- River Otters ing from all other mustelids in Wisconsin. The hard palate extends posteriorly, well beyond “Particular points of Lutra proper are the presence the last molars (but never as far as 15 mm). of perfect claws, in comparison with their absence The upper carnassial (P 4/)is broad and non- or rudimentary condition in some other genera, and shearing, and the posterior upper molar is the lack of special dilatation of the tail.” — Elliot subquadrate or rhombic, not triangular as in Coues, 1877. Taxidea or dumb bell shaped as in mustelines. The interorbital constriction is narrow 1762. Lutra Brisson. Regnum animale..., ed. 2, p. (narrower than the rostrum), the zygomata 13. Type Mustela lutra Linnaeus. Brisson’s slender, but the zygomatic width is broad and names are not consistently binominal, and are the infraorbital canal is large and oval. The regarded as non-Linnaean. rounded braincase is not much elevated above 1772. Lutra Bruennich. Zoologiae Fundamenta... p. the short rostrum, so that the entire skull 34. Type Mustela lutra Linnaeus. seems flattened, and the sagittal crest is only 1843. Lontra Gray. Ann. Mag. Nat. Hist., ser. 1, 11: weakly developed. The ear structure is much 118. Type Lutra canadensis Schreber. Van as in skunks (Mephitis), and the auditory bul- Zyll de Jong (1972) advocated use of this name lae are similarly flattened. for the North American river otters. A mathematical appraisal by Van Zyll de Jong (1972) found American and Eurasian Hall and Kelson (1959) and Hall (1981) otters in separate groupings. Recent work on provide a detailed generic synonymy under mitochondrial cytochrome b sequences also the name Lutra. suggests their divergence, and in an old-fash- The river otters, usually only called “ot- ioned morphological comparison they also ter,” are elongate aquatic mustelids having differ (albeit only slightly). The American riv- weak scent glands; short legs with webbed, er otters have Gray’s name Lontra. pentadactyl feet; short and dense brownish fur; and a cylindrical body tapered at both ends. The short-haired, furred tail is thick at Lontra canadensis (Schreber) the base; the neck is as thick as the head. River Otter

Lontra canadensis canadensis (Schreber)

1776. Mustela lutra canadensis Schreber. Die Saugthiere... Theil 3(Heft 18), pl. 126. Type from eastern Canada (= Quebec). The trivial name canadensis is available for the river otters, but the trivial name lutra applies to Euro- pean otters. 1823. Lutra canadensis: Sabine. In Franklin, Narra- tive of a journey to the shores of the Polar Sea... 1819-22. P. 653. 1972. Lontra canadensis: Van Zyll de Jong. A systematic review of the Nearctic and Neotropical river otters (Genus Lutra, Mustelidae, Carnivo- ra). Royal Ontario Museum Life Science Contr., Sketch of river otter. C.B. Cory, 1912. 80: 1-104. See also Koepfli and Wayne (1998).

414 THE WILD MAMMALS OF WISCONSIN The name Lontra means otter in Latin, club shaped. Its product of width and length and the word canadensis is Latinized, mean- exceeding 400 mm indicates sexual maturity ing the animal was discovered in Canada. (Friley, 1949). The chromosomes are 2N=38; Otter was a European word for the closely the fundamental number 62 (Wurster-Hill and related European member of the genus. It is Benirschke, 1968). derived from the ancient Sanskrit udra, mean- The fur is a rich brown or chestnut ing “an aquatic mammal.” In northern Europe brown, with lighter grayish brown underfur. d is pronounced like t, or the other way The underparts are paler brown with paler around, and hence the words “otter” or Lutra, underfur, which when wet shows abundant- and even Lontra. The Wisconsin race associ- ly. Around the cheeks the fur is light brown ated with the Great Lakes region in Michigan or tan, with prominent vibrissae. The eyes are and Wisconsin, usually dwells in the smaller black. There are five toes on each foot which lakes and marshes and along rivers and show in the tracks. In juvenile otters the color streams in that region. Otters have extended seems brighter, with orange-brown on the their range into the Mississippi drainage fol- back. Wisconsin otters are much darker than lowing that great river southward. my one Florida specimen. Jackson (1961) Description. See generic account above. suggests there are two molts; possibly there Baker (1983) aptly refers to the shape of the is one gradual molt through summer, which skull constricted in dorsal view as “nearly hour- is difficult to appraise in small samples. glass shaped.” The dental formula is unique Melquist and Dronkert (1987) mention for Wisconsin mustelids, 36 teeth. No other molts in spring and autumn. carnivore’s body is tapered at both ends, with Males are a little larger than females, a stout, furred (i.e., short-haired velvety) tail. about 4 percent or more (Hooper and Osten- The ears are rounded and quite short; they son, 1949) and as much as 17 percent heavi- can be closed under water. The eyes are small er (Melquist and Dronkert, 1987). Hoffmeis- and set forward in the head. In addition to ter (1989) gave the measurements in milli- the cranial characters mentioned above, the meters of a single otter from Peoria, Illinois, coronoid process of the lower jaw is vertical as 1,120 mm total length, 396 mm tail, and and quite high, the lower molars are basined 144 mm hind foot length (45 inches, 16, and for crushing, and the mandible occasionally 6). The interorbital breadth was 24.25 mm, (with age) locks into the squamosal as in and the maxillary toothrow was 37.1 mm. Taxidea or Gulo. The baculum is long and Baker gave the external measurements in mm for Michigan otters of males as 950-1,300, 320-445, 110-125, and ear length 10-25. His weights of males ranged up to 25 pounds (11.3 kg). For three adult Wisconsin otters external measurements are as follows: male 1152, 415, 116, 16; females 1025, 375, — , —; 960, 415, 116, 16. Mean cranial measurements (and extremes) for four adult males and three females were, respectively, greatest length of skull 109.4 (108-113) 110.9 (108-113), cranial breadth 60.0 (55-65) 60.2 (56.4-62.2); zygomatic breadth 74.3 (66- 78.6) 72.9 (66-78.6); interorbital breadth 26.8 (21.8-27.1) 24.8 (24.4-25.5); maxillary Skull of Lontra (= Lutra) canadensis. tooth-row 36.8 (35.5-37.6) 37.5 (36.5-38.5).

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 415 Dental Formula. I 3/3, C 1/1, P 4/3, M 1/2 = 36. Occasionally there is a supernumerary premolar. Geographic Distribution. The river otter formerly occurred throughout the entire state in lacustrine or riparian habitats, and owing to legal protection (with restricted trapping) it has reoccupied its former range in remote places. It may be found in any Wisconsin county, except probably Waukesha and Milwau- kee counties, but the animal is most abundant in the northern lake country, also ranging along large and medium rivers to the

Maps showing geographic distribution of Lontra canadensis in Wisconsin and North America.

416 THE WILD MAMMALS OF WISCONSIN southward. The river otter is not known on them. Depressed fur markets will allow them any islands in the Green Bay area, but re- perhaps to become numerous enough for portedly otters do occur in the Apostles (Jack- general enjoyment. It is a joy to watch a fam- son, 1961). I have seen river otters swimming ily of otters sporting in the water like seals. in the sea from San Juan Island to Brown The chief benefits of otters are that they have Island in the state of Washington, and no valuable fur and are wonderful, graceful aquat- doubt they occasionally cross Green Bay to ic creatures to watch in nature. The few neg- the islands there. ative aspects of otters are killing ducks and Status. The river otter is fairly abundant fishes (although few game fishes are eaten, in Wisconsin, enough so that the Wisconsin Knudsen and Hale, 1968). The leading coun- Department of Natural Resources allows lim- ties for harvests (1994-1999) were Oneida ited trapping in some counties. After protec- (84), Sawyer (68), Price (66) and Taylor (63). tive legislation was enacted in 1915, the ot- The northwest district has the highest num- ter steadily increased in numbers (Jackson, ber. For example, in 1995, the leading coun- 1961). The species reoccupied most of its ties were Price (118) and Barron (78) (Dhuey former range, which retreated to northern and Kohn, 1995) and Chippewa Native Amer- counties in the late 1800’s. icans took another dozen. Males are taken Bluett (1984) reported the harvest data more frequently (751:451). There were no from 1970-1983, finding the low of 483 in records of otters taken along the Illinois bor- 1970-71. Numbers increased until 1975-76, der except in Grant County, and none from when 853 otters were taken. the Door Peninsula. Nearly 70 percent were The annual harvests in Wisconsin have trapped in Conibear traps. About 25 % were fluctuated by about 1,000 otters more or less trapped in beaver sets. and pelt prices varied from $82 to $24. No In 1994-1995, the harvest was high more than two (and usually one) otters were (4,615 pelts) with an average price of $48.76. permitted per season. The harvest in 1994- The total value approached a quarter of a mil- 95 was 1,816 registered otters, with an aver- lion dollars. Some other recent statistics pro- age pelt value of $48.76, somewhat less than vided by Dhuey (personal correspondence) the previous year when 1,130 were harvest- show these interesting results or the seasons ed for an average price of $57.30 (Dhuey of 1996-1997 and 1995-1996, respectively: and Kohn, 1994, 1995). Recent pelt price 1,368 and 1,233 pelts [fewer], $45.66 and was as low as $12.50 (in 1992), which seems 44.47 [about as high], and [only] $62,463 and low for fur that this author regards as the most 54,832. The only reason I can see that possi- luxurious of any. In the winter of 1999-2000, bly explains such results is that the number of the harvest was one of the greatest ever, trappers, the number of otters, or both, was 2,178 otters (Dhuey, Kohn and Olson, 2000). high. In 1999, 2,178 otters and 187 bobcats In Upper Michigan the river otter was were harvested (Dhuey, see Kohn et al. 1998. never in much peril, although the numbers Habitat. The habitat requirements are had declined. It has been protected since water in which to swim, stream banks above 1925, with complete protection until 1939. the water suitable for denning, and sufficient Today it is fairly abundant (Baker, 1983). aquatic foods, such as fishes. Polluted streams There are some records in Illinois (Hoffmeis- and human harassment can cause otters to ter, 1989), which was another vast region abandon these sites. Otters usually live remote where the otter was perhaps eradicated and from people unless protected. The otter thrives was reintroduced in the 1990’s. in deep bodies of water, as well as shallow Urbanization definitely is a threat to ot- marshes, and they move along streams, search- ters, but recent wetlands protection will help ing the banks and pools for prey.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 417 In Illinois one burrow had no nesting ma- Otters are swift in the water and can terial inside; the entrance was a few feet above outswim fish. They also corner them or sur- the water edge. There were a few sticks in the prise them by leaping on them from overhead burrow. Liers (1958) mentioned nest-lining* in banks. Doubtless some fishes eaten are carri- the natal den. In Stevens Point, an adult fe- on. According to Bluett (1984) his studies in male with young used a short burrow in the Michigan, Wisconsin, and Minnesota reveal bank of an offshore island. This den was prob- that otters feed primarily on rough fish and ably a temporary den or outlier, and was sited do not affect the number of trout (except for a few feet above the water level (and water an otter resident in a fish hatchery). He de- table). It was probably made by muskrats.* tails foods (percentages) of Wisconsin otters Foods. Fishes make up most of the diet as fishes (77 %), crayfish (13%), and frogs in Minnesota (Liers, 1951), Michigan, and (6%). Game fishes did not exceed 22%. Suck- Wisconsin (Knudsen and Hale, 1968; Alex- ers, minnows, carp, and mud minnows com- ander, 1977; Field, 1970; Greer, 1955; La- prised 48 percent of the total diet. In Melquist gler and Ostensen, 1942; Ryder, 1955). and Dronkert (1987) the frequency of occur- However, some panfishes and game fishes rence (percentages) in the diet of river otters are eaten. Many fishes eaten are suckers and was listed as fish 70-99%, crustaceans 3-75%, other kinds of the less appreciated “forage” insects 1-46%, amphibians, 1-20%, birds 1- fishes. Lagler and Ostenson (1942) found 13%, and small mammals, 1-25%. The crus- about 36 percent of the food was forage fish- taceans are usually crayfishes, the birds wa- es, and about 23 percent were game and pan terfowl), and amphibians frogs. fishes. Sometimes feeding on overcrowded Other food reports include crayfish, panfish is actually a benefit for them, allow- aquatic insects, clams, mink, frogs, and musk- ing these fish to grow larger. As Baker (1983) rat. On land or snow otters catch, in winter, stated, “the ecologist takes the attitude that meadow voles, red-backed voles, and even an there should be fish enough for human and occasional snowshoe hare (Field, 1970). Berry otter alike . . . There certainly was abundance seeds (Miller, 1937) and remains of birds (Lag- of these prized fish in the early days when lar and Ostenson, 1942) were eaten. Otters otter were probably more numerous along refused to eat deer carcasses set on the banks streams than they are today.” to tempt them. A definitive study on the diet of the otter Reproduction. Otters begin reproducing is provided for the upper Midwest by Knuts- usually in their second year. Males mate fe- en and Hale (1968). The primary foods were males promiscuously. Copulation has been fishes and crayfishes. This was true no mat- observed in water, lasting up to 25 minutes ter the season Whether one considered the (Melquist and Dronkert, 1987). After parturi- occurrence or volume, the story was the same tion in late March and early April (Ostenson (see Table 17). The preferred fish species were and Gross, 1940) and after mating and con- suckers, mudminnows, minnows, and sunfish- ception, new embryos form. Blastocysts are es. Trout were surprisingly low in volume or delayed from implanting until the following occurrence. The study included otters from year. Gestation may even last 12 months Wisconsin, Michigan and Minnesota. Rare (Hamilton and Eadie, 1964). True gestation foods were a leech, fairyshrimp, Gordiacean is about 60-63 days. I have a lactating Wis- worm, clam, and two turtles. consin specimen from April 27. Litter size is about 2-3, up to six (but there are only four abdominal mammae). In Wisconsin the mean * Otters may appropriate woodchuck burrows for natal dens. I have observed as a huge otter covered himself with fallen leaves on my litter size is 2.4 + 0.5 (N=14). Mean number river bank before falling to sleep. of corpora lutea was 2.3 (N= 42). The new-

418 THE WILD MAMMALS OF WISCONSIN born are reportedly pink but furred, but edge, following streams or riverbanks, but is Melquist and Dronkert (1987) report some are more oval in marshes. Females in winter move pigmented black, with conspicuous vibrissae, very little, covering only 4.8 mi 2. Males seem eyes closed, toothless, weighing about 275 g to have larger home ranges than females. Lit- (9.6 oz) each. Melquist and Dronkert (1987) tle information is available on home range and give the weight of the neonate as 120-160 density of young, yearlings, mature adults, grams (4.2-5.6 oz). The eyes open by 35 days, males, females, or of the extent of movements and by 40 days the young are playful. They in water. Dispersal occurs when the otter is emerge in two months from the den and are 12-13 months old, but not all yearlings dis- weaned by three months. The family may stay perse significant distances. In 16 months one together for 7-8 months. By 75 days, usually otter used 88 different dens and resting sites in June, the young go swimming with their (Melquist and Dronkert, 1987). Beaver dens mother. Some males do not breed until they both active and inactive were often used (38 are five years of age. Longevity in the wild percent). On snow a male may traverse dis- may be as long as 10 to 15 years, but 13-19 tances of up to 3 miles (4.8 km) (Field, 1970). years in captivity (Crandall, 1964). In winter two otters may occupy the same Aging can be accomplished by counting den; one may even occupy the same den with cementum layers (Stephenson, 1977), matu- beaver (Baker, 1983). rity (ossification) of the baculum, and tooth Densities are never high. Jenkins (1949) wear. Some old otters have worn down or examined 212 otters from the Upper Penin- missing teeth. sula of Michigan. One third was yearlings, Mortality. Humans harvest otters by trap- mostly males. ping, but many otters are road-killed near bridg- Remarks. River otters (even the adults) es on highways. Pollutants may be important spend considerable time playing. They slide causes of mortality because in polluted waters down mud and snow banks, wrestle, and tum- the otters feed at the top of aquatic food chains. ble about. Usually the adult males are soli- Otters might be eaten by various predators, tary, as are the females when they have no e.g., coyotes and wolves, especially when out family. Otters are nocturnal, but they are of- on land or on snow-covered land. In central ten seen by day. Wisconsin one occasionally sees the otter’s trail Additional Natural History. Lariviere and striking off through a marsh, easily recognized Walton (1998) published a review on the riv- by the sledding track of its round body and er otter. Harris (1968) published a study on short legs made moving over the snow. Coy- the natural history of the Lutrinae, including otes range through the same marshes. American species. Three ticks have been collected on these Geographic variation. There is only one mammals; it is surprising that the water does race in Wisconsin and Upper Michigan. not drown them or wash them off (Jackson, Hoffmeister (1989) assigned all Illinois speci- 1961). Flukes and nematodes are internal parasites. Melquist and Dronkert (1987) list canine Table Car-17Car-17. Volumes of foods found in Wisconsin ot- distemper, feline panleukopenia, hepatitis, jaun- ter digestive tracts. Percent volume Cubic centimeters. After dice, pneumonia, and tuberculosis. These have Knudsen and Hale (1968). never proved much of a problem in Wisconsin. Stomachs 6,137 Intestines 1,726 Home Range and Density. Melquist and Hornocker (1979) have studied populations Fishes 87% 47% Crayfishes 4 46 and abundance by trapping records, tracking Insects Trace 5 on snow, and radio-telemetry. Obviously, Frogs 7 1 home range shape depends on the water’s Mammals 1 2

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 419 mens to L. c. lataxina Cuvier, following Van seem related to the primitive, Asian ferret Zyll de Jong. Jackson (1961) assigned all Wis- badgers (Melogale). The number of mustelid consin otters to L. c. canadensis. This is not a teeth has been reduced from 38 to 34. In typical Jackson-Hoffmeister phenomenon with connection with its more predatory habits, the state boundary exactly dividing the two Taxidea has not broadened the cheek teeth races, because there is a hiatus in the range. as much as omnivorous Eurasian Meles or Most Illinois otters are in river drainages that Asian Arctonyx and (smaller) Mydaus, which drain southward in Illinois. Most Wisconsin are comparable in body size and appearance. otters are boreal, from habitats northward of Taxidea has a broad, wedge-shaped head that the tension zone or within it. In the last few is much less globular, and as a consequence decades L. c. canadensis has been spreading lacks the sagittal crest seen in Meles. The southward in Wisconsin. Along huge rivers they somewhat sectorial upper carnassial is large may make long dispersals. Hoffmeister (1989) and robust, but possesses the inner accessory recorded a recent otter killed by an automo- cusp found in true badgers. The coronoid bile in Whiteside County, 2 mi. S Albany, Illi- process of the dentary is not recurved poste- nois, and less than half a mile from the Missis- riorly (Long and Killingley, 1983). sippi River. That otter may well have been an Badgers are of medium size for carnivores emigrant from northern L. c. canadensis. in North America, about the size of a stout, Specimens examined. Total, 17. Adams, short-legged, and medium-sized dog or raccoon Bayfield, Crawford, Marathon, Marinette, Procyon. The forepaws are huge and the fore- Oneida, Portage*, Vilas, Waupaca, Wood claws are as long as 5 cm. The innermost claw counties. At the Kansas Museum Nat. Hist., is relatively thin. The toes are partially webbed are two specimens not seen (I have seen xerox for strength and digging. In the collagen su- copies of the skulls and labels). 10 mi. W Rhine- perstructures associated with the phalanges lander and 12 mi. N Rhinelander, Oneida Co. Pacinian corpuscles are found, indicating a Other records are after Balliett and Taft (1978): sense of feel not discovered in any other mam- Chippewa and Clark counties. Not plotted. mal (Long and Killingley, 1983). The large nictitating membrane doubtless protects the eyeball from dirt (Long, 1972; Long and Killing- Genus Taxidea Waterhouse ley, 1983). As in all badgers, the Taxidea have North American Badgers anal scent glands (with pervasive but not powerful odor). The head and dorsum, at least the 1778. Ursus, in part, of Schreber. Die Saugthiere, head and nape of the neck are conspicuously 3:250. marked with advertisement patterns, comprised 1784. Meles, variety americanus of Boddaert. Elen- in Taxidea of a median, white dorsal stripe, chus Animalium, 1:136. and black “badges” on white cheeks. 1823. Taxus labradoricus Say. Long’s expedition... 1:261, 369. Not Taxus gulo of Tiedemann. 1838. Taxidea Waterhouse. Proc. Zool. Soc. Lon- don, p. 154. Type species is Ursus taxus Schreber.

The North American badgers are an off- shoot of ancient, long-extinct badgers, and

* A photo of an adult was in Stevens Point Journal, Thurs., March 24, 1977. Sketch of badger. C. Long and C. Killingley, 1983.

420 THE WILD MAMMALS OF WISCONSIN Taxidea taxus (Schreber) mixed with gray, brown, dull ochraceous, tan, North American Badger and whitish hairs. The cheek teeth may not be as robust as in the T. t. taxus, but the “The strange fact that some of the pioneers in shape of the skull is similar (Long, 1972). Wisconsin were called “badgers” encouraged a The badger is easily distinguished from fondness of the people for this animal and fostered other carnivores in Wisconsin by the triangu- an appreciation of its beneficial habits. It became an lar upper carnassial or the upper molar, which endearing symbol”. — Charles A. Long and Carl A. is also triangular. The auditory bullae are well Killingley, 1983. inflated, never low as in skunks or the river otter. The low, broad shape of the posterior Synonomy given above under the genus. part of the skull, with its wedge shape seen There are two size groups in North Amer- from above, the huge forefeet with enormous ica, the larger northern badgers and the small- claws (see Figure), the shaggy fur and short er southern badger (Long, 1972). Three sub- tail, and the pattern of median white stripe species comprise the northern form, and the and black badges on the cheeks, low ears southern is a subspecies marked by a long defended with hairs, and even the hooked and median stripe often to the base of the tail. The twisted tip of the baculum are quite distinc- dorsal pelage is variable in color and seems tive. Chromosomes are 2N = 32. grayer in the eastern parts of its range where The bold median stripe on the head and it intergrades with the grayish race of the Great neck of the Wisconsin badger is not known Plains. There is much microgeographic varia- to extend onto the back, contrary to some of tion, as in bears, but the races recognized the pictures on early roadmaps of Wisconsin. (Long, 1972) are well differentiated except Usually this is a trait of the southernmost sub- where confused with minor family and local species, which is smaller than our badger. The variations or by intergradation in broad zones crown of the head and the badges on the of hybridization. Cranially, the Wisconsin race cheeks are black or dark brown. The cheeks Taxidea taxus jacksoni Schantz resembles the are usually whitish or creamy buff, but black nominate race Taxidea taxus taxus. fur hides the black eyes. The low ears are edged with whitish hairs. The feet are blackish or dark brown. The shag- Taxidea taxus jacksoni Schantz gy fur of the back, which may be erected in a threatening display, has intermixed colors of 1945. Taxidea taxus jacksoni Schantz. J.Mammal., 26:431. Type from 4 mi. E Milton, Wisconsin.

Both Taxidea and taxus mean badger. This race was named after Wisconsin’s own Hartley H. T. Jackson, a scientist in the Unit- ed States Biological Surveys, U. S. National Museum, by Viola Schantz. Description. See the account of the genus above. The badger is the only fossorial carnivore in Wisconsin, interesting in its adaptations of form, function, and behavior. This northern race T. t. jacksoni resembles in coat pattern and size the nominate race. It is darker, more brownish and even blackish inter- Skull of Taxidea taxus.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 421 black, brown, buff, gray, yellowish tan, and reddish chestnut. The coarse guard hairs are tipped with white or pale buff. The color varies in appearance owing in part to how the fur is ruffled, and how much of the lighter underfur (sometimes called wool) is exposed. The paler, buffier juvenile molts to adult pelage, and there is an annual molt thereafter (Long, 1975). New hair seems to appear first in the dorsal stripe, extending posteriorly, the molt band widens until the molt is completed. Prob- ably in Wisconsin it occurs from May to June, perhaps July (Long and Killingley, 1983).

Maps showing geographic distribution of Taxidea taxus in Wisconsin and North America.

422 THE WILD MAMMALS OF WISCONSIN Weights in adult males approach 25 mining excavations in southwest Wisconsin, pounds (11.3 kg), and about 40 lbs. in a Wis- and due to the courageous defensive demean- consin captive from Magnussen’s Fur Farm or of this independent denizen. Even though (Long and Killingley, 1983). Males are signif- thriving the state does not wish to permit the icantly larger than females (Long and Killing- state symbol to be hunted or trapped, and ley, 1983; Messick, 1987). An old male from the aesthetic values of this interesting preda- Portage County measured 838-125-72, with tor maintain its status as protected in all sea- condylobasal length 130.0, zygomatic (84.0) sons. The animal is a furbearer, depending and cranial (79.3) breadths, and maxillary on the market and fashion often quite valu- tooth-row 42.3 mm. able, recently $100 per pelt, and this has led Dental Formula. The formula is I 3/3, C to some poaching. The fur is coarse, although 1/1, P 3/3, M 1/2 = 34. Dental abnormali- God valued it (of the Eurasian badger of ties and their incidence have been described course) enough to tent the tabernacle of the by Long and Long (1965), Long and Killing- Israelites (Long, 1987). It is hardly valued for ley (1983), and Messick (1987). They usually coats, except by some people who wear them are subnumerary or supernumerary premo- to University of Wisconsin—Madison football lars or caused by injury. games (the mascot is “Bucky Badger”). The Geographic Distribution. Reportedly T. fur is valued higher when style demands it, t. jacksoni occurs in all counties of the Up- and often it is used to tip furs or as trim on per Peninsula of Michigan (Baker, 1983), and garments. In the past the badger fur was fa- may be expected in any county in Wisconsin. mous for art, paint, and shaving brushes, but Badgers are uncommon in the forested coun- Meles was the badger usually utilized in this ties, but thrive in prairies, including the hilly way (Long and Killingley, 1983). “goat prairies” of western Wisconsin, and The Eurasian badger was persecuted in many wet marshes, riparian meadows, and Britain, “badgered” by vicious dogs in the pit sandy tall-grass and short-grass prairies of and by other tortuous methods. Even in Amer- Wisconsin. They seem restricted northward ica Taxidea was occasionally pitted against beyond the canal at Sturgeon Bay, unknown dogs. A badger in a steel trap is an epitome to my knowledge on the Door Peninsula, and of cruelty because of its indominatible nature have not established themselves on any known (see Long and Killingley, 1983). The badger island in Lake Superior or Lake Michigan. The is a symbol of humanity’s conversion to com- geographic race has somehow crossed into passion for animals. Ontario above the Superior-Huron Waterway The belief that the badger-holes break the Connection (Baker, 1983) where it in peril legs of horses and cows leads to some exter- (Long, 1972; Long and Killingley, 1983; mination, mostly undeserved (I know of not a Long, 1992). single instance of such injury to livestock, nor Status. At this time the badger is doing any person thrown from a horse that stum- very well in Wisconsin, expanding its range bled in a badger hole). Badger holes occasion- where it was previously unknown or had been ally hasten erosion of dikes and roadways, but extirpated, and increasing in numbers (Leroy what excavator could raise subsoil and in sev- et al., 1975; Long and Killingley, 1983; Long, eral ways contribute to soil formation? 1992; and Wydeven, Wiedenhoeft, and Dhu- Habitats. Badgers mostly live in open ey, 1999). The badger is a beneficial mam- prairies, usually well drained or hilly so that mal helping farmers in their struggle against the burrows are above the water table. There- rodent and rabbit pests, and is also the state fore, badgers often build dens on dikes and animal and state symbol. Pioneers were called railroad rights-of-way that are levated, grassy, badgers due to their living in dugouts and their and weedy. In savannas and meadows one

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 423 can occasionally encounter a badger among foods. See table Car-18. No studies have been trees and undergrowth. They are often found made on Wisconsin badger foods, but Errington in association with their prey, normally ro- (1937) studied them in northwestern Iowa. dents of one kind or another. In Minnesota Another fine study was made in Iowa by Snead and western Wisconsin, the prey species is and Hendrickson (1942). Mostly the badger usually Geomys bursarius; the habitat there- feeds on rodents and other small vertebrates, fore is grasslands, pastures, roadsides, and occasionally on plant foods when in season or sandy hillsides. In central Wisconsin mice, when the badger is hungry. It eats many insects ground birds, rabbits, insects and other prey and other invertebrates, carrion, and when avail- are opportunistically taken, and the habitats able nestlings and eggs of ground-nesting birds. are more variable. The soils where badgers live Lampe (1976, and other works) presented a are friable, and usually sandy. Badgers range great deal of information on food physiology around the southern shore dunes of Lake Su- and predation of Minnesota badgers. perior on the Upper Peninsula of Michigan. Usually the badger eats up its kill, but Except for the natal den or the winter den, occasionally caches rabbits and rodents in a badger diggings tend to be located with their burrow. The prey is captured underground, prey. A badger may excavate a den for only a but occasionally the badger emerges to run day or two, use and move on to a new burrow down prey. regularly thereafter. There may be as many as three natal dens used (Lindzey, 1976). Where a badger lives as long as a few weeks, when prey is abundant, the burrow system usually has a central chamber with one to four tunnels to the surface and several blind passages. A burrow may have a nest (grasses, but sometimes the nest chamber is bare) two or three feet, even six feet down, at the end of an entrance tunnel as short as eight feet and as long as 30 feet. The entrance is often oval or ellipsoidal because the badger is so short-legged and stout of body, and the diameter varies from 19-24 cm. The entrance may be plugged with sand and a mound of sand emitted from the excavation marks the site. Natal dens (see Lindzey, 1976) have twice as large mounds, and the tunnel bifurcates so that two tunnel branches are present. These come back together again, and there are usually several blind pockets and passages. There may be a second entrance. Scats are found in short pockets usually covered over with dirt. Sometimes there are two resting chambers. The branching tunnels allow badgers to pass by one another going and coming. Foods. The badger is a predator, mostly Development of skull in Taxidea, showing fusion of on injurious rodents of one kind or another. It sutures and broadening of the skull with age. Long, details is an opportunist with an incredible variety of in 1975.

424 THE WILD MAMMALS OF WISCONSIN Reproduction. Badgers pair in summer There is an unconfirmed record (Schwartz and or autumn, about the same time the males Schwartz, 1981) of a litter of seven. Some become reproductive. The females probably young-of-the-year females become pregnant. do not come into estrus until July as a rule, In Wisconsin, there have been no studies on although some copulate as early as May and badger reproduction. June (Messick, 1987) and heat lasts through Skulls can be arranged in age classes (see August, depending on the altitude and lati- Fig.) as juveniles (milk teeth present), young tude somewhat. Copulation was described by (with basioccipital-basisphenoid suture open), Campbell and Clark (see Long and Killingley, subadults (basioccipital-basisphenoid suture 1983), with the male holding the female’s fused, but nasal sutures unfused, and teeth neck with its jaws. Copulation lasted 21 min- lack wear), adults (nasal sutures fused, teeth utes. The successful male was observed to fight worn), and old adults (cranial sutures fused, with another, and they emitted musty scents. sagittal crest evident, and postorbital process- Badgers wound one another frequently es well developed) (see figure, also Long and during the breeding season (Todd, 1980). Long, 1975). Messick (1987) reviews numer- Messick (1987) and Messick and Hornocker ous aging techniques, bacular size and cemen- (1981) reported ovulation from July to August. tum rings (see Crowe and Strickland, 1975) After fertilization the zygote develops to being particularly accurate. He also compares the blastocyst stage, which in a stage of quies- the age classes by mean body weights. In cap- cence have been recovered from many bad- tivity badgers may live 20 years. gers up until the time of implantation in the In old badgers the squamosal fossa so spring. This delayed implantation (Wright, encircles the condyloid process the lower jaw 1966; 1969; Hamlett, 1932; and others) al- is locked fast (Long, 1965, Long and Killing- lows the young to develop and be born when ley, 1983). This accomplishes several advan- the mother has emerged from her winter sleep, tageous adaptations: preserving the fulcrum rodents are abundant, and the growing sea- of crushing, self-sharpening the canine teeth, son has begun. The implantation occurs in late and allowing the lower canines, as they shear, February in Idaho (Messick and Hornocker, to brace the upper ones against breaking off. 1981) and Montana (Wright, 1966). Thus, Mortality. Aside from man the badger has delayed development was about 6 1/2 months. few enemies. People often shoot and trap Gestation lasted about five to six weeks. Lac- them, and automobiles take a toll, especially tation lasts until June, maybe a little longer in of males. Coyotes and dogs could conceiv- some females. Young are brought forth in mid- ably kill one caught above ground. Young March to early April plus or minus a week or badgers probably are killed by raptors and so, and lactation was observed in May. several carnivores. Adult badgers are so fierce Badger young are born furred, but blind and intimidating they are seldom attacked and helpless. Their eyes open at four to six even by bears. weeks of age. Weaning occurs when the Parasites and disease doubtless affect young are about half grown, the young are badger mortality greatly. Long and Killingley taught to hunt when about two/thirds grown (1983) list the parasites which are flukes, tape- and weaning is past. The mother brings food worms, nematodes (eight species), ticks (six items to the natal den for the young. The male species), fleas (nine species), and a malloph- does not help rear the young. agan louse. Western badgers often test posi- Litter size is usually two to four young tive for plague and are no doubt susecptible (Messick, 1987; Long and Killingley 1983). to some other diseases of carnivores. Appar- Observed mean litter size varied from 2.2- ently some juveniles starve (Messick, 1987) 2.74, with standard deviation about 1.0. and old badgers may do so (Long and Killing-

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 425 ley, 1983). Messick (1987) presents detailed on fat, which is often their only source of studies of mortality and life tables. Most bad- energy in winter. They do regularly enter pe- gers die in their first year, and nearly as many riods of torpor, observed to last about 29 yearlings die (Messick and Hornocker, 1981; hours, when the heart reduces its rate by half, Messick, 1987). Long and Killingley (1983) and the body temperature falls about 9 de- pointed out that young are dependent on their grees C. Fats accumulate in the blood, and in mother for nearly a year, including six months delayed implantation. Table Car-18. The great variety of badger foods eaten Home Range and Density. Badgers are by an opportunistic forager and hunter. After Long and Kill- solitary except when accompanying their ingley (1983). young or mating. They are territorial between males, which fight over females, and the Insectivores Shrews Blarina, Sorex mother defends her den, young and feeding Carnivores territory. There have been no population stud- Coyote Canis latrans pups ies on badgers in Wisconsin, but a radio- Skunks Mephitis, Spilogale tracked badger in Minnesota wandered 850 Rodents hectares, in summer, only 53 in the fall, and Marmots Marmota monax, flaviventris* Chipmunks Tamias, Eutamias in winter only 2 hectares (five acres) (Sargeant Ground Squirrels Spermophilus, Ammospermophilus and Warner, 1972). Lampe (1976) found a Prairie Dog Cynomys* home range of 1700 hectares in summer for Red Squirrel Tamiasciurus another female. He found seasonal variation Gophers Geomys, Thomomys*, Pappogeomys* in home range size, diminishing toward win- K-rats Dipodomys* Harvest Mice Reithrodontomys ter (Lampe, 1980; Lampe and Sovada, 1981). Deer Mice Peromyscus When food is abundant badgers seldom wan- Other rodents Onychomys*, Ondatra, Mus, Rattus, Microtus, der, except the breeding males. Lindzey Lagurus*, Synaptomys, Zapus (1978) found wandering confined to 270-627 Lagomorpha hectares. Messick and Hornocker (1981) de- Rabbits Sylvilagus Hares Lepus termined home range to vary with age. Adults Birds had home ranges only 0.62.4 km2. In winter Grebe Colymbus the animals seldom wander about during their Bank Swallows Riparia period of winter sleep. Male home ranges Ducks Anas (eggs) overlap female home ranges. Messick and Pheasant Phasianus (eggs) Reptiles Hornocker (1981) found that in areas of abun- Lizards Phrynosoma* Uta*, Cnemidophorous, Crotaphytus* dance the density is as high as 3-5/ km2. Snakes Thamnophis, Crotalus, Pituophis Remarks. Badgers are extremely fosso- Turtles Chrysemys rial, especially for such a large mammal. They Amphibia have been observed to dig through an asphalt Frogs Rana, Toads Bufo Fish Carp Carpio road, and can bury themselves on the spot as Arthropods you watch them. Lampe (1976) carefully de- Insects Acridae, Gryllidae, Carabidae, Apidae, Vespidae, scribed their digging behavior. Often bisect- Formicidae, caterpillers homopterans ing a tunnel prior to localizing the prey to Arachnids Spiders, Scorpions* one side or the other, the badger was suc- Chilopoda Centipedes Plants cessful 73 percent of the time. Behavior of Corn, oats wild and captive badgers is reviewed in Long Other plants Flowers, grasses, Solanum, and Killingley (1983). Carrion Badgers do not hibernate and may be Fish, ravens, chickens, sheep, coyotes, hares, pheasants active on warm days in winter. They do lay * Not found in Wisconsin

426 THE WILD MAMMALS OF WISCONSIN many ways the badger resembles a hiberna- cats such as lions, house cats, and lynxes are tor (Harlow, 1981). The Germans call this arranged. Usually the cheetahs are arranged phenomenon “Winterschlaf” or wintersleep, in a separate genus. In such an arrangement in the European badger Meles. the Canada lynx, European lynx, perhaps the Additional natural history. Long and Kill- Spanish lynx, and the bobcat usually are ar- ingley (1983) reviewed the biology of the ranged in the subgenus Lynx. Some taxono- North American badger. mists do not use subgenera, and tend to rec- Geographic Variation. The badger shows ognize several cat genera, including Lynx. In no geographic variation in Wisconsin, but grades North America, at least, the lynx and bobcat south and west. The subspecies is weakly dif- are distinctive from the house cat or moun- ferentiated, more nearly black than other rac- tain lion. A number of recent workers using es. This dark appearance may result from natu- skeletal measurements, principal component ral selection owing to millennia of dwelling in analysis, antibodies, and karyotypes have bol- wet prairies and marshes, riparian meadows, stered the argument for the old classification and floodplains near boreal and post-glacial (genus Lynx) in opposition to those prefer- woodlands of the North Woods Region, even ring Felis (Lynx) (for example, see Hoffmeis- though it dwells in Wisconsin on the highland ter, 1989). One may ask cui bono? It is merely prairies. Specimens just southward of the Wis- a crux criticorum, with little biological sig- consin boundary in Illinois are also dark, indi- nificance, to either separate Lynx as a genus cating an affinity to the endemic, dark lake-states or subgenus. subspecies, instead of to some recent prairie invader into Illinois. Numerous badgers now found in Douglas County may be prairie immi- Genus Lynx Kerr grants with paler pelage, but those examined Lynx and Bobcat occurring in nearby northern Minnesota were of the dark jacksoni race (Long, 1964a). Many changes of names are merely opinions Specimens examined. Total, 29. Buffa- of taxonomists, and a great many names de- lo, Dane, Dodge, Iowa, Jefferson, Ke- rive from those who make taxonomic and waunee, Marathon, Marinette, Marquette, prestigious compilations. Changes should re- Portage, Sauk, Waushara, Winnebago, sult from a comprehensive study of specimens Wood counties. of a taxonomic group, and presentation of Other records (Long, 1972): Rock Co., quantitative, i.e., sound reasons for making 4 mi. E Milton, Vilas Co., Mamie Lake. Wal- worth Co., Delavan and Emerald Grove.

Family FELIDAE Gray Cats

The felids have short, arched skulls with elongate canines and an evolutionary reduction of molars above and below. Behind the enormous shearing carnassials little remains with which to chew. The cats stab, slice, and swallow. The claws are retractile. The genus Felis is to some authorities a Sketch of the Lynx preying on a snowshoe hare. Lloyd huge taxonomic umbrella under which diverse Sanford in W. J. Hamilton, Jr.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 427 the name changes. Examples of less than rig- portionally larger and the supporting orbital orous argument include some opinions on the fascia hypertrophied, and the great tendons problematic scientific names of the Canada of the toes of the Canada lynx are distinctive lynx and North American bobcat. in size. Other papers focus attention on the Kurten and Rausch (1959) were ambiva- distinctness of Lynx in the patterns of princi- lent, saying that the statistical difference be- pal component analysis (Werdelin, 1981), tween European and American Lynx was molecular immunological distance (Collier and small. Perhaps the two cats were even con- O’Brien, 1985), and chromosomal patterns specific. Then both would be called Lynx lynx. (Kratochvil, 1982). Therefore, I believe there Because they apparently are closely related, is reason for recognizing Lynx as a full genus. the inclusion of European Lynx with Ameri- can Lynx at the generic level must follow, but inclusion of the Caracal (Felis caracal = Cara- Lynx canadensis Kerr cal caracal, from Africa and Asia) with them Canada lynx becomes problematic. Including diverse cats in the genus Felis with Lynx is likewise problem- Lynx canadensis canadensis (Kerr) atic, for many interesting distinctions are obscured. Modern classifications of cats today 1792. Lynx canadensis Kerr. The animal kingdom, boggle the mind, with one recognizing two felid 1:157. Type, eastern Canada, Quebec. genera, another four, another 12, another 14, 1758. Felis lynx Linnaeus. Syst. naturae, ed. 10. and one with 17 genera*. 1966. Felis (Lynx) canadensis: Van Zyll de Jong. Van Gelder (1977) mentioned a [possi- Canadian J. Zool., 44:499. ble] Lynx-Felis hybrid, suggesting both kinds belonged to one genus, but Hall (1981) rec- The generic name Lynx means “lamp- ognized Lynx as a distinct genus. Jones et al. sight,” taken usually to refer to the bright eyes. (1992) also used Lynx. Several recent work- I suspect the Canada lynx was observed at ers have used the genus Lynx. Lynx closely night in lantern light, which created a bright resembles Felis in cranial characters, with eye shine. The name canadensis means from some trivial difference of the shape of the Canada. Mammalogists call this animal the premaxillaries. Of importance is a dental dif- Canada lynx, but some people call it the lynx ference in the number of premolars, only two cat or wild cat. pairs above in Lynx and three above in Felis. Description. The Canada lynx is a large Different numbers of teeth are often used in cat (a meter in length, about 30 pounds weight classifying distinctive genera of mammals. ( = 13.6 kg), bob-tailed, with enormous ball- The tail is short in Lynx, which seems like feet. Its large tracks (with claw prints usu- trivial on the face of it, but taken as one of a ally lacking, claws retracted) may be confused suite of cold-climate adaptations, such as the with those of a mountain lion. One specimen’s large feet for walking on the snow, the tufted forepaw was larger than the hind foot of a ears, and the shaggy ruff on each jowl, the large male wolf. The paw measured 110 X short tail takes on significance. (If Old World 120 mm, not including the spreading mar- caracals belong to Lynx, then some lynx at ginal hairs. The pointed, erect ears are least would have longer tails.) To this I add my adorned with long, prominent tufts of hair, observations that the eyes in Lynx are pro- and the jowls or cheeks are margined with a mane-like fringe or ruff framing the face. The face is characterized by huge eyes, and the * Compare that taxonomic hodge-podge of the closely resembling cats with another Carnivoran family with truly diverse genera, such belly may have a typical sag or paunchiness as the Mustelidae. seen in some of the big cats. The hind foot

428 THE WILD MAMMALS OF WISCONSIN exceeds 200 mm in length, but the entire foot dorsally, are tipped with black and there is a of a digitigrade cat is not entirely on the subterminal band of white. The underfur is ground. The rear end of the lynx body is some- cinnamon brown. The upper parts are griz- what elevated compared to the front end. The zled white, black, brown, tawny, and gray. hind foot has but four toes. There is a (fifth) The head is the same color but there is a toe, elevated, called a dew claw, on each fore- profusion of white-tipped hairs. The ears are foot. There is one pair of abdominal and one white inside and brown outside, and the tufts pair inguinal mammae. The baculum is short- are black. The eyelids are white. The ruff or ened, probably vestigial. fringe around the face is a mixture of brown, The skull resembles in shape that of the black, and white hairs. The sides of the body mountain lion, domestic cat, or bobcat (espe- are somewhat ochraceous; the underparts cially the bobcat). It is short-faced, with elon- whitish or buffy. The molt begins in late gate canines, short tooth row (with only two spring when the fur is generally brown. The premolars); the carnassial premolar is elon- color becomes more grayish as the guard gate and sectorial; and the upper molar is tiny, hairs grow. somewhat vestigial. Males are slightly larger on average than The zygomata are extremely bowed but females. The body length may be as long as the postorbital process is not as robust as in 1,000 mm, and the weight as much as 39 the bobcat. The dentary is nearly straight ven- pounds ( = 17.6 kg). The body weight usually trally as far as the root of the lower canine. varies from 15-25 pounds (6.8-11.3 kg). Full The distinguishing characters of the lynx growth is attained in the third year (Quinn skull, and these are not always distinct, are and Gardner, 1984). Greatest length of the the presphenoid shape and certain cranial skull is approximately 140-150 mm, and zy- foramina (see figure of the base of the crani- gomatic breadth varies from 90-100 mm. um). The presphenoid is broad posteriorly but Dental Formula. I 3/3, C 1/1, P 2/2, M greatly constricted anteriorly. It resembles a 1/1 = 28. tiny spade. The anterior condyloid foramen Geographic range. Today the Canada and the posterior lacerate foramen (also called lynx may not permanently live and breed in the jugal foramen) have separate openings Wisconsin. Possibly they are all wanderers near the auditory bulla. The bobcat’s presphe- from Minnesota, dispersing southward and noid is narrow, and the two foramina share a eastward into this state. Jackson (1961) re- common fossa. viewed the records and reports of Wisconsin The color overall is brownish gray. The lynx up to that date, and found originally they dorsal guard hairs, which are darkest mid- occurred from the western counties to the Door Peninsula, southward sporadically as far as Dane and Milwaukee counties and southern Jefferson County. Specimens and sightings known to Jackson (1961) were documented since the turn of the century until about 1960. There is no evidence proving the lynx might have persisted in Wisconsin until recently, but it is not easy to document extirpation (Doll et al., 1957). Some sightings in recent years are reported now and then in the western and even in the southern coun- Base of braincase showing presphenoid shape and anterior ties, especially curious was the roadkill from condyloid foramen. Bobcat is to the left. the hilly, southwestern woodlands.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 429 In Upper Michigan, the lynx seemingly ed for sport or trapped commercially. In its existed until at least 1976 (Long, 1976). Dr. behavior, so wonderfully adapted to the bo- Phillip Humphrey told me at that time there real forests and to the deep winter snow, and were probably no lynxes left in Upper Michi- so majestic its appearance, the Canada lynx gan. There has been a natural spread of lynx is one of the most beloved of Wisconsin mam- from Ontario between Lakes Superior and mals. It is unfortunate that the lynx is so rare. Huron, where the lynx extended its geograph- Never common, and in some periods likely ic range into Michigan, into regions where they eradicated, the lynx periodically reappears. probably had been in peril or had vanished. When any do reinvade the state, they might Michigan lynx apparently also dispersed into be shot mistakenly or illegally by hunters. Oth- northern Wisconsin (Emlen 1954, Iron Coun- er lynx die naturally from disease, perhaps, or ty, personal corr. to H. H. T. Jackson; and old age, or they may be killed on the high- Doll et al., 1957 Washburn County, January ways. Although this mammal is a protected 22, 1956). Lynx immigration into both Wis- wild animal (Nat. Resources 10 Wisconsin consin and Upper Michigan may have been Administrative Code) in Wisconsin, so long as primarily from Minnesota, as it seems today. cars drive down the highways and hunters kill Status. This large and interesting cat is cats in the wild there is little hope for natural beneficial to humankind, seldom preying on re-establishment of this rare cat. Another lim- livestock (although it may feed on carrion) and iting factor is the possible competition between preying often on rabbits, snowshoe hares, and the lynx and the bobcat (which is difficult to mice. The fur is valuable. In Canada, the har- measure and a word to use with caution). Of- vest varies from about 10,000 to a maximum ten the greatest barrier to an animals’ invasion of 50,000 each year, a pelt might sell for is a closely related animal. The point of in- $600 (as in 1984), and a lynx fur coat may gress for many Canada lynx is Douglas Coun- sell for $20,000. Pelts of protected species ty, where bobcats now seem abundant. may not be sold in Wisconsin. If it were legal Thiel (1987) reviewed the status of Wis- to harvest the Canada lynx, it would be hunt- consin lynx, reporting sporadic and cyclic occurrences mostly following periods when Lynx cycles are at peak abundance in Cana- da. Inasmuch as the bobcat has widely replaced the lynx, in Wisconsin, even taking over the boreal habitats, and now ranging deep into Canada, the interrelations between these two similar cats should be studied to understand if the lynx forfeits or competes poorly with the bobcat. Neither species is harmful to humankind; indeed it would be a joy to observe them both. The Canada lynx has been protected as an Endangered Species since 1973, and as a protected wild animal in Wis- consin since 1998. Recently (March 2000) the U. S. Fish and Wildlife Service listed the Canada lynx as “Threatened” under the Rare and Endangered Species Act, and it would again be listed as endangered if suddenly it Geographic range in North America for the Lynx seemed endangered in a significant part of its canadensis. geographic range. This implies surveillance,

430 THE WILD MAMMALS OF WISCONSIN and recovery plans are in effect. Another lim- shoe hares, and opportunistically on other iting factor is the change in habitat caused by prey when hares decline in numbers. Hares forestry practices in the northern counties; comprise about 60 percent of the winter diet, habitat for lynx in the Great Lakes region is 40 percent of the summer foods. Lynx will ranked as “marginal.” Prey densities (i.e., cache hares, but not always do they return to Lepus americanus) may not sustain lynx pop- the cache to eat them. These data (Car-19) ulations, and cutting out softwoods and plant- from Quinn and Parker (1987) are based on ing of aspen may be detrimental to lynx. [How- earlier reports of Brand and Keith (1979), ever, near Ely, Minnesota, new aspen growth Nellis and Keith (1968), Van Zyll de Jong is common range for lynx.] Trapping and (1966), and Stewart (1973). hunting are not considered of much impor- Animals other than hares, mice and tance in conservation and recovery. That grouse that are eaten by lynx on occasison seems to write off much in recovery for this include red squirrels, flying squirrels Glau- species in Wisconsin. However, perhaps the comys sabrinus, beaver, red foxes, striped state of Wisconsin can face the challenge, skunks, porcupines, masked shrews, proba- even transplanting lynx here and there and bly raccoons, and some other uncommon replanting some softwoods, which is some- foods (Youngman 1975; Saunders 1963; thing of an investment anyhow. I am uncon- Nellis and Keith 1968; Nellis et al. 1972). vinced that our resources in the vast tama- The Canada lynx rarely feeds on plant mate- rack swamps and marshlands, and the densi- rials, and occasionally kills a fawn or moose ty of snowshoe hares and present abundance calf (but seldom kills adult deer). No doubt of whitetails for food are inadequate for lynx. they scavenge on adult deer carcasses. Habitat. The Canada lynx inhabits heavily Reproduction. The mating time in Lynx forested areas, especially the boreal conifer- is brief, from mid-March to early April (Quinn ous forests with heavy snowfall. Lynx ca- and Parker, 1987). If food is abundant, young nadensis will inhabit and have inhabited farm females may breed, but most lynx breed as country, but only when it is interrupted with yearlings or later. Whether ovulation is in- forests (Quinn and Parker, 1987). Parker duced or not is debated. Van Zyll de Jong (1981) found the habitat preference was strict, (1963) suggests the ovulation is spontaneous. avoidance of hardwoods and favoring of se- The gestation period is nine weeks; the young ral conifer and climax conifer woodlands. The are born furred and blind in late May and ear- use by lynx of coniferous forest increases in ly June. The eyes open in about 9-12 days, summer. Usually the lynx live with the snow- perhaps 14 days (Quinn and Parker, 1987). shoe hares (Lepus americanus). The Cana- At birth each kit weighs about 200 g. Soft da lynx avoids dense alder (Alnus spp.) swales, pelage replaces the woolly natal coat in 60 according to Parker (1981). days and the adult pelage is assumed in nine According to Jackson (1961) the natal months. The kittens are weaned by 12 weeks, den is in a hollow tree, stump, or log, and but stay with the female into the winter, when sometimes is sited under fallen timber. The the young disperse (Saunders 1963, 1964). nest consists of leaves, bark, and other vege- The rates of pregnancy and litter size depend tal material pawed and trampled into a form a great deal on the abundance of Lepus. Lit- by the female. Sometimes a lynx hides in a ter size is about 2-3, reportedly as high as den, but usually it prefers to rest crouching 4.6 in peak years, and ranges from 1-6. Lon- on top of a rock or log where it can watch for gevity is between 11-12 years, possibly long- prey or enemies. er (see Crandall, 1964). Food. Lynx prey selectively, and almost Mortality. Aside from hunting and trap- entirely when they are abundant, on snow- ping, there is little information about preda-

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 431 tors of Lynx. The “kits” might die from star- climbs well, swims well, and of course it is well vation or fall prey to large carnivores and large adapted to leaping on snow. The great paws owls. Jackson (1961) lists five species of fleas will support double the weight of the smaller and two kinds of tapeworms (Taenia) as par- paws of the bobcat before breaking through a asites. Nematodes include hookworm, Tox- crust of snow (Quinn and Parker, 1987). ascaris, Toxocara , and Troglostrongylus Lynx Population Cycle. The now famous (Quinn and Parker, 1987). A nematode Cyl- multiannual population cycles seen in lynx icospirura may be found in cysts in the stom- populations, involving browse, prey, and pred- ach wall, as many as 70 worms per cyst. Their ator, closely corresponds to the period of their effects are unknown. Lynx can suffer rabies principal prey, the snowshoe hare (Elton and and feline distemper. Even the male adult Nicholson 1942, Nellis et al. 1972, Keith, Lynx may kill lynx kittens that they find, a 1963; Gunderson, 1978; Mech, 1980; and behavior also seen in other cats. others). Fur records show the period has been Home Range and Density. The home fairly regular since 1820. Professor Lloyd range, greater in males than in females, was Keith, of the UW-Madison, and his research about 16-20 km2, but varied enormously from associates, have made numerous long-range 12 to 243 kilometers square (Saunders 1963, studies on this cyclic fluctuation. In Canada, Berry 1973, Brand et al. 1976, Mech 1980, the periods of the predator Lynx and the Parker et al. 1983). Mech’s findings (1980) snowshoe hare cycle on average every 9-10 suggest, in Minnesota, that the home range years. Brand et al. (1979) and Nellis et al. is much greater than previously thought. (1972) suggest lynx numbers fall because of Ranges varied from 51 to 122 km2 for fe- post partum starvation of kittens caused by a males and 145 to 243 km2 for males. scarcity of hares; perhaps fewer female Lynx Density estimates are extremely variable. conceive. Population densities varied from 2.3 To this variation is the added complication per 100 sq. km. (winter 1966-67) to 10 per that many data are from far away, i.e., Rus- 100 sq. km. (1971-2). sia, Alaska, the Maritime Provinces, and so The cycle, long considered a classic exam- forth. More information for Michigan and ple of a predator-prey interaction, is actually a Wisconsin is needed. The lynx varied in den- browse-prey-predatory cycle. The snowshoe sity from nearly 0 to 10/100 km square, dur- hares increase in numbers tremendously eating ing several winters in Alberta (Brand and Kei- up their forage, and begin to feed on inferior th 1979). Movements on a nightly basis in and indigestible foods. This forage quality sends Minnesota averaged 3.2 km; the cats showed the hare populations into decline, and the lynx overlapping home range (Mech, 1980) par- predation combines with malnutrition to crash ticularly of females. the hare population. The scarcity of hares sends The male is solitary, associating with the the lynx into decline, not so much by starvation female only during the brief mating period. as by causing smaller litters and fewer pregnan- There seems to be scent marking of territories cies of female lynx. The immigrant lynxes I have with urine. The Canada lynx is essentially noc- dissected in winter (1992) were heavily layered turnal (Baker 1983; Quinn and Parker, 1987). with fat, even on the crown of the skull, and Hunting follows three techniques, namely fol- contained ample fat deposits within the abdo- lowing snowshoe hare trails, concentrating men. The intestinal tract was stuffed, and the predation within small areas of snowshoe hare fur in good condition. activity, and crouching [lurking] beside snow- After the hares decline to low densities, the shoe hares’ trails. A mother with young may forage plants begin to recover their abundance. advance through vegetation to flush snow- The lynx had meanwhile become fewer by lower shoes, which the cats all pursue. The lynx reproduction, natural mortality, and some emi-

432 THE WILD MAMMALS OF WISCONSIN gration. In the absence of their predators and Sect. 31, T29N, R18W 1 1992. Vilas Co. with improving forage the hares increase in num- Vic. Woodruff, 1 UW 1972. bers again, eventually to reach another peak. The Other records: Long 1970. Portage Co., timing is about 9-10 years concurrently over S Bancroft. Vernon Co., Veroqua. 1966 or enormous areas in numerous states and prov- 1967. UWSP- Exhibits 1. Jackson Co. City inces (Bryant 1981, Keith 1974, Keith et al. Point 1966 or 1967. County records by Thiel 1984, Quinn and Parker 1987). (1987) include 1963 (Douglas); 1965? Thiel (1987) compared these peak fluctu- (Pierce), 1965 Green Lake, 1965 (Vernon), ations with museum specimen occurrences, in a gap until 1971 (Trempealeau), 1972 (Trem- Michigan, Wisconsin, and Minnesota. The pealeau), 1972 (Oneida), 1972 (Price), 1973 specimens in these 3 states seem to result from (Iron), 1974 (Marinette). Reportedly the Rusk peak populations in Canada. Wydeven (1993) and Ashland specimens are bobcats. reported 3 carcasses for 1992, a year of pre- dicted dispersals. One invasion was in 1971- 1972, and another was in 1992. Certainly all Lynx rufus (Schreber) specimens collected of Lynx canadensis should Bobcat be preserved in permanent museum collections. Thiel (1987) mentioned failures in preserving “The bobcat is a shy and furtive in its behavior and specimens of lynx. Some preserved as lynx are very seldom seen. [It] does a service for the in fact specimens of bobcats (Wydeven and Iwen, ranchman in keeping down the numbers of Rabbits personal correspondence). and small rodents. — H. E. Anthony. The first bona Geographic Variation. There is a single fide field guide to mammals, Field Book of North subspecies in Wisconsin and Upper Michigan American Mammals, 1928. Specimens examined. Total, 5. Burnette Co. Town of Anderson 1 1992. Marinette Co. 1 Univ. Wisconsin Campus at Marinette. Chippewa Co., near Edson 1. St. Croix Co.

Table Car-19. Foods of Lynx. Quinn and Parker.

Winter in Alberta: Snowshoe hares 35-90 % Mice and voles 4-28 % Squirrels 9-12 % Grouse 2-6 % Miscellaneous 13 % Northwest Territories* 33 %/60 % 19%/ 7% 11% /1% 3 %/ 7 % Remainder miscellaneous. Ontario** 63 % /70 % 0 % / 4% 0 % / 3 % 6 % / 5% Remainder miscellaneous. *Summer and winter **Fall and winter Sketch of Bobcat. By John A. Litvaitis. June 2000.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 433 1777. Felis rufa (Schreber). Die Saeugthiere... Theil forefoot, bobcat tracks show only the impres- 3, p. 95, plate 109b. Type from New York. sions of the lower four toes, two to each side 1817. Lynx rufus: Rafinesque. Amer. Monthly for each forefoot. The tracks may show the Mag., 2: 46. impressions of claws, depending whether or not they are retracted. The binomen means red cat, and Lynx The dorsal fur is dappled brownish or refers to a lamp-shine, probably from the great tan, intermixed with reddish or grayish but eyes in reflected light. Bobcat refers to the less grayish than in the lynx. There is an in- “bobbed” (short) tail. termixture of buff and some black, which is somewhat intensified on the head. The underparts and inside surfaces of the legs are Lynx rufus superiorensis white with an admixture of dark spots. The Peterson and Downing dorsal surfaces of the ears are black with a prominent white spot visible on each. The 1952. Lynx rufus superiorensis Peterson and Down- ears are tufted with black. The tail is white ing. Contr. Royal Ontario Mus. Zool. and Pa- below, with a black dorsal band on the tip. leontol., 33: 1. Type from McIntyre Twsp. near This black and white tip differs from the all Port Arthur, Ontario. black tip in the Canada lynx. There are several obscure bands closer to the rump. The The bobcat, wildcat, or in earlier litera- white and black marks on the ears and tail ture catamount, unfortunately also goes un- may signal the young to follow the mother der two scientific names. Van Gelder (1977) in heavy undergrowth (Rolley, 1987). and others have considered Lynx as insepa- The total length varies from 82-95 cm rable from the genus Felis. Others (e.g., Wer- (to 37 inches) for males, and 73-85 cm in delen, 1981) recognize Lynx. females. The tail is only 12-16 cm in length. Description. This cat is twice as large as The weights range up to 15.4 kg (34 pounds), a housecat and usually (but not always) small- but average 12.7 kg (28 pounds). Females er than the Canada lynx, which the bobcat vary up to 15.4 kg (34 pounds), averaging closely resembles. The bobcat has large eyes 6.8 kg (15 pounds). A male bobcat killed in set forward, dense, soft dappled fur, and its 1970, near Greenwood, Clark County was ear tufts are relatively smaller than those of 51 inches (1.29 m) in length and weighed 38 the Canada lynx. pounds (17.2 kg). I have a two-by-two col- As in most felids the skull is short, the ored slide of a bobcat shot in Ashland Coun- cranium arched, and the canines curve downward and hook somewhat posteriorly. Some facial hairs are long and hang from the jowls like a beard. The cheek teeth are highly evolved for shear, with elongate carnassials, and a reduced upper molar. The distinguishing characters that separate bobcat from lynx are given in the account of the Canada lynx. There are four mammae, two inguinal and two abdominal. The karyotype is 2N=38 (Hsu and Benirschke, 1970). The os penis is a small vestigial ossicle in the glans. The partially webbed forepaws are small with retractable claws. Although a dew claw is present on each L. rufus. Note presphenoid.

434 THE WILD MAMMALS OF WISCONSIN ty in December 1982, which weighed 42.5 ing each annual harvest and regulating hunt- lbs (photo by Larry Gregg). Lariviere and ing and trapping. The estimated population size Walton (1997) list measurements as 86.9 (to varies according to the harvest of the previous 125.2), 78.6 (61-109); tail length (14.8, 13.7; season. No hunting or trapping is permitted hind foot length 17 (13.0-22.3), 15.5 (12.5- southward of Highway 64. State-wide, and 19), ear length 66, 63 for males and females, especially in the central savannas and sands, respectively. Greatest length of skull and zy- the bobcats are less common than 30 years gomatic breadth were 129.3 + 5.8, 120.7 + ago. Public attitudes in Wisconsin are more 5.7 and 89.8 + 5, 84.2 + 4.4. enlightened today; and the bobcat is not shot Dental formula. I 3/3, C 1/1, P 2/2, M as “vermin” as it once was. Possibly there are 1/1 =28. There are 30 teeth in the house 2,000-3,000 bobcats living in the state, most- cat and mountain lion, usually assigned to the ly in the North where the similar Canada lynx genus Felis, and more than 30 in other Wis- ought to be re-established. Seldom are bob- consin Carnivores. cats seen in the south where some thriving Geographic range. Originally the bobcat populations ought to occur. Therefore, no ranged throughout the state, especially the hunting is permitted there. Hunting and trap- bottomlands and rocky coulees of southern ping both are legally carried out in northern Wisconsin, and the swamps and forestlands counties, but the populations seem to be hold- of northern Wisconsin. There the range over- ing their own. lapped that of the Canada lynx. Jackson In 1988, the total harvest was 165 ani- (1961) concluded the former range included, mals; most of these were shot by hunters us- in suitable habitats, the entire state. There ing dogs, and 46 were trapped. The counties was ecological replacement northward, be- with the largest harvests were Marinette (24), cause the Canada lynx was better adapted to Douglas (20), Oneida (16), Price (13), and the snow and to feeding on larger prey. Rusk (13). Dhuey and Kohn (1995) made a Today, the primary Wisconsin range of Life Table on aged bobcats, and the estimat- the bobcat is northern Wisconsin, particular- ed population size was seen to be increasing ly the forested northwest corner. The bobcat somewhat, from lows in the early 1980s of is seldom encountered today south of the Ten- about 1,500 animals to over 1700 in the early sion Zone. Habitats for it are plentiful in the 1990s, based on the Model. coulee country along the Mississippi, where Kohn and Ashbrenner (1995) report the the undergrowth of prickly ash and brambles population of bobcats has steadily increased should provide a haven for their renewal. Re- from 1,300 in 1988 to 1,950 in 1995. Dhu- sults of a DNR survey of records from 1980 ey and Kohn (1995) reported 169 were killed to 1990, show some occurrences south of in 1994, and 160 in 1993. The pelt price Highway 64. The bobcat is absent today from (about $36) was lower than in 1993 ($45) the heavily populated regions of south-cen- and much lower than in the 1980s. In 1997, tral and southeastern Wisconsin. It occurred the pelt price was $44.75, but the harvest in recent years in the Jordan Swamp, the was low. The harvest for 1999 was 187. The pines near the Tomorrow River, and near flesh and fur of the bobcat in Wisconsin Junction City, all in Portage County, central amount to little value, although individual pelts Wisconsin. bring high prices. Status. Historically, the bobcat may have Bobcat pelts sold in the late 1970s ap- followed a 10-year population cycle of period- proached 150,000 in Canada and over ic abundance according to Jackson (1961). 80,000 in the United States (Obbard et al., Today populations are managed by Wisconsin quoted in Rolley, 1987). The U.S. prohibited Department of Natural Resources by assess- exports of bobcat pelts in 1981, and the cur-

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 435 rent hostility by animal rights groups toward the fur trade diminished the number of pelts taken and sold. Furs usually are used for taxi- dermy mounts and wall rugs. Regulation of the bobcat harvest includes seasons, possession limits, restrictions on hunters and trappers, export quotas, and potentially a possible emergency closure of the season. It is easy to over-harvest mammal species with low reproductive rates, and wildlife managers must set harvests conservatively. Sometimes closing of seasons may be necessary. The bobcat is a beneficial mammal, for it feeds on rab-

Maps showing geographic distribution of Lynx rufus in Wisconsin and North America.

436 THE WILD MAMMALS OF WISCONSIN bits, mice and injurious animals. Beyond that, Reproduction. Breeding time (generally, in it is important in our legends and lore, beau- winter continuing into early spring) may be in- tiful to observe in nature, but is seldom seen. fluenced by photoperiod, latitude, climate, nu- Habitat. The bobcat prefers habitat with trition, and the age-composition of the popula- abundant prey, dense vegetation in bottom- tion. Ovulation may not be induced as previ- lands or mixed conifer forests, and rocky sites ously thought. It occurs usually from February for natal dens. Deep snow, human settle- through March. Gestation is about 62 days (Mc- ments, and deforestation probably are adverse Cord and Cordoza, 1982). If there are two lit- factors. Dense-canopy forests are avoided ters, obviously the bobcats may breed in other (Rolley 1987). In the northern parts of the months. In Wisconsin, only one litter is known range, bobcats prefer stands of Norway spruce to be produced each year. Litter size varies from (Picea abies), mixed hardwoods, and eastern 1-6, with averages of 2.5 to 3.9. Yearlings pro- hemlock (Tsuga canadensis). Snowshoe duce smaller litters. Kittens are born furred with hares, which are often abundant in cedar eyes closed, and they open in 3-11 days. The swamps and black spruce, attract bobcats to weight of the newborn is about 283-340 g. these snowy habitats. These dense woodlands Young are suckled for about two months. Young provide habitats with less wind. accompany the mother, hunting together at 3 In Wisconsin, Lovallo et al. (1993) and months, and disperse prior to the breeding sea- Lovallo (1993) studied bobcats at natal dens son, usually the following spring (McCord and in Douglas County. They were found in a Cordoza, 1982). Lovallo (1993) found dens in sedge meadow with Spiraea, Salix, and Al- May in Wisconsin. However, he suspected some nus in scattered clumps; near Picea and Thuja breeding later. The life span seldom reaches stands with intermixed paper birch (Betula 12 years, but one reportedly lived 32 years in papyrifera); and in cover types identified by captivity (Jones, 1982). 1986 infrared photos he listed as follows: 1. Mortality. Man and his dogs, starvation, Young pines, 2. Lowland conifer (Thuja, disease and injuries comprise the main caus- Larix, Picea), 3. Unforested grasses and sedges of mortality. Natural mortality is rather low es, 4. Oak burn, 5. Mixed burn (Quercus, (Rolley, 1987). The bobcat suffers rabies, cat Pinus), 6. Upland deciduous (Populus, Acer, scratch fever and pneumonia (Lariviere and Tilia), 7. Young aspen, 8. Upland conifer, 9. Walton, 1997). Dubey et al. (1987) reported Mixed forest on uplands, and 10. Lowland lethal toxoplasmosis. Parasites observed in alder thicket (Fraxinus, Alnus). Breeding bob- Wisconsin by Lovallo et al. (1993) included cats preferred upland deciduous forest, and Felicola, Levineia felis, and Isospora felis. also used lowland conifer extensively. Other parasites include helminths, protozo- Dens included an abandoned beaver ans, mites, lice, fleas, and ticks. lodge with the bobcat’s entrance above water Home Range and Density. Bobcats of the and a log-brush pile in a clear cut area. Natal same sex seem to avoid one another (Rolley dens are dry, hard to find, and usually in a 1987). Home ranges of female bobcats hard- remote area. Females often move their young ly overlap (suggesting territorial defense), but to outlier dens. Resting sites include rocks, males tend to wander in and out of territo- brush, windfalls, and hollow trees (Lariviere ries. Home ranges vary greatly (0.6 to 201 2 and Walton, 1997). km ) and are influenced by sex, density of Foods. Snowshoe hares, cottontails, and bobcats, and prey availability. Scent marks the white-tailed deer (mostly sickly or carrion but territories (with urine, feces and anal secre- also an occasional adult) are the chief foods. tions) (Lariviere and Walton, 1997). Other foods include mice, squirrels, opossums, Lovallo (1993) estimated home range in birds, and snakes. Douglas County, by use of radio monitoring.

TAXONOMIC ACCOUNTS / ORDER CARNIVORA 437 Fourteen resident bobcats were located at least Specimens examined. Total, 4. Forest 50 times each. The smallest home range for Co. No specific locality, 1. Oneida Co. 10 2 a female was 7.18 km2 (9.06 km by use of mi. W Rhinelander, 1. Near Rhinelander, 1. the harmonic mean). The largest male home Portage Co. Junction City, 1. Other records 2 range was 184.27 km (149.43 by use of the (Balliett and Taft, 1978): Shawano Co. No harmonic mean). Males are likely to wander, specific locality. Stevens Point Jour. Feb 28, and they spent longer times doing so judging 1990, Mead Wildlife Area, probably Wood by their activity patterns. The average home Co. and Milwaukee Jour., near Greenwood, 2 range for females was 29 km , and for males Clark Co., 1970. 2 96.9 or 85.8 km . McCord and Cordoza 2 (1982) report densities of 0.04 per km . A male captured 10 June 1992, on the Order ARTIODACTYLA St. Croix River, moved during the following Even-toed Ungulates 14 days northeasterly until it was lost from radio contact on 24 June, several miles east Hoofed mammals, usually large (more than of Solon Springs. The minimum distance tra- 100 pounds adult weight), of which the body versed between these two points was 40.8 weight is borne between the third and fourth km. Bobcats have larger home ranges in the toes, usually with lateral vestigial toes (called northern part of their range, where resourc- dew hoofs), and the astragalus has articular es are sparse and habitat quality is poor. In surfaces (and is said to resemble a double- Minnesota they are forced to wander more pulley) at either end. Pigs are bunodont, ar- extensively and for longer distances accord- tiodactyls selenodont (with crescentic cusps). ing to Fuller et al. (1985). Bovidae, with keratinous-bonecore horns dif- Geographic Variation. This species has fer from deer (Cervidae), in which males have been reported to have about a dozen geo- branched, deciduous antlers. Advanced artio- graphic races in North America, although the dactyls have a ruminant (four-chambered) validity of some is questionable. In Wisconsin stomach. In Wisconsin, there are domestic no geographic variation was noted for the artiodactyls, including pigs, cattle, sheep, bobcat. Possibly the prairie race, from Illinois, goats, llamas, alpacas, and re-established Bi- formerly occurred in southern Wisconsin prai- son. The only wild, self-sufficient artiodactyls ries (see Jackson, 1961), but in recent years in Wisconsin are the white-tailed deer and the bobcats reported in southern Wisconsin prob- moose. The domesticated horse Equus ca- ably dispersed there from northern Wiscon- ballus is in a different order, Perissodactyla. sin (from the race Lynx rufus superiorensis). That situation may soon change, as bobcats now ranging widely in Illinois may invade the southern counties. The presence of one race of bobcat in Wisconsin and a different one in Illinois is a different example of a Jackson-Hoffmeister phenomenon, because there was a hiatus between the two populations caused by extensive clearing of woodlands and other land uses. Hoffmeister and Jackson both were probably correct, in this instance, but the boundary between the races cannot be ascertained. Sketch of hoofs of Artiodactyla. Caton.

438 THE WILD MAMMALS OF WISCONSIN Key to Artiodactyla in Wisconsin Family CERVIDAE Gray Deer, American Elk (or Wapiti), and 1. Horny sheaths have bone cores, which Moose are extensions of the frontal bones, horns present in both sexes, nasal chamber The deer family is characterized by antlers overlain with bone ...... Bovidae (usually seen only in males, a year old or old- Bison bison (extirpated, er), which are usually branched extensions, now semi-domesticated); domestic smooth and polished, of the frontal bones. cow, Bos taurus; sheep, Ovis aries; When growing they are blunt and rounded at and goat, Capra hircus the tips of the tines, and covered by fur and 1’ Horns lacking, nasal cavity overlain with vascularized skin, not at all resembling the bone, teeth bunodont ...... keratinous horn of the Bovidae. Like bovids, ...... Suidae, domestic pig the deer have selenodont and hypsodont Sus scrofa cheek teeth, a ruminant stomach (four-cham- 1" Horns lacking, but antlers present in adult bered) and cloven hooves (i.e., two large toes males, antorbital vacuity or fossa opens with hooves on each foot, with the body into nasal chamber, teeth selenodont .. weight on the legs transmitted to the evenly ...... Cervidae...2 divided toes), which is shown in an accompa- 2 Adult pelage spotted on dorsum, found nying sketch. Large antlers found buried along in southeast Wisconsin ...... streams are often the extinct Cervus elaphus ...... Introduced deer…..2a elaphus; all elk antlers resemble those in the 2a Antlers palmate, mane lacking ...... following sketch...... Fallow deer, Dama dama 2b Antlers not palmate, mane present ...... Sika deer, Cervus nippon 2’ Adult pelage never spotted ...... 3 3 Antlers palmate, pendulous “bell” up to 7 seven inches length suspended from throat, muzzle large ...... Moose, Alces alces. 3’ Antlers not palmate, “bell” lacking, muzzle delicate and conical ...... 4 4 Posterior narial cavity not completely divided by vomer, upper canines usually present but vestigial, antlers elongate. Small herd confined to NW Wisconsin and also maintained on approximately 100 game farms in Wisconsin ...... Introduced American Elk, Cervus elaphus nelsoni 4’ Posterior narial cavity divided by vomer septum, canines lacking above, antlers having a curved main beam from which tines rise vertically .... White-tailed deer, Odocoileus virginianus Sketch of antlers of elk from Caton.

TAXONOMIC ACCOUNTS / ORDER ARTIODACTYLA 439 Genus Odocoileus Rafinesque Odocoileus virginianus (Zimmermann) North American Deer White-tailed Deer

Odocoileus Rafinesque, 1832, is not the old- Odocoileus virginianus borealis Miller est generic name for the American deer, but the older name Dama is currently applied to 1900. Odocoileus americanus borealis Miller. Bull. the European fallow deer. Adhering to no- New York State Mus. Nat. Hist., 8:83. Type menclatural priority would replace our Amer- from Bucksport, Maine ican Odocoileus with the name of a popular 1902. Dama v[irginiana]. borealis: J.A. Allen. Bull. and much-studied European deer, causing Amer. Mus. Nat. Hist., 16:20 unnecessary confusion. The International 1919. Cariacus wisconsinensis Berlitz. Wis. Cons., Commission on Zoological Nomenclature 1:1, for Dama virginiana Zimmermann, 1780, used its plenary powers to conserve Odocoile- Geog. Geschichte, 2:24. us for American deer (Bull. Zool. Nomencl., 1965. Odocoileus virginianus borealis: Kellogg. In 17: 267-275, 1960). Taylor’s The deer of North America.

Referring to the selenodont cheek teeth, the name Odocoileus means hollowed out teeth. The name virginianus agrees with Odocoileus, whereas virginiana does not grammatically, and it means from Virginia. Usually called whitetail or white-tailed deer, the species has also been called the Virginia deer. The word deer is Old Norse for beast. Adult males are “bucks”, females “does”, young of the year “fawns”, and year-old deer “yearlings.” Foot of whitetail. Spencer Fullerton Baird. Sometimes a huge and stately old male is called a “stag,” as in E. T. Seton’s “Trail of the Sandhill Stag”. More often the word stag is applied to Cervus or Rangifer. Description. In deer, the hooves are paired on each foot, and on each forefoot a pair of interesting vestigial “dew hooves” persist, one on either side of the foot. These are inherited toes from the ancestral artiodactyls; during evolution they became smaller. Whitetails are easily recognized owing to their size (90-130 kg), slender legs, distinctive antlers in males, long ears and slender muzzle, and the conspicuous tail, which is brownish above and pure white below. The skull of the whitetail and other deer shows intricate “fractal” sutures bordering the parietals and supraoccipital (Long, 1985; Geographic distribution of Odocoileus virginianus in Long and Long, 1992), which evolved com- North America. plexity makes the suture much stronger to

440 THE WILD MAMMALS OF WISCONSIN bear the great weight and forces transmitted The molt, including both hair loss and by the antlers to the skull. A large antorbital its replacement to gray, winter fur, commenc- fossa opens into the nasal scroll bones anteri- es in late summer or early autumn. The light- or to each orbit. Pronghorns (Antilocapra er, red summer coat appears in late spring americana) also have such a fossa. (depending on climate and latitude, late May From the western mule or black-tailed to early June). In Central Wisconsin, I have deer, Odocoileus hemionis, the whitetail dif- regularly observed deer with patches of gray fers in the color and shape of the tail (which falling out and red fur appearing in early May, is flag-like, and not blackish and slender). Also, and gray fur was nearly completely replaced the antlers branch differently. In O. virgin- in October. Some deer show patches of sum- ianus there are usually short tines projecting mer red lingering about the rump as late as upwards from a main beam, whereas in O. mid-October. The gray brown pelage replac- hemionus the antlers branch dichotomously es the red throughout September. Fawns are for two or more branchings. reddish-brown or reddish-tan, with white The upper parts of the whitetail are gray- spots dappling the dorsum. These disappear brown in the winter, darkest mid-dorsally, and at 3-4 months of age (Sauer, 1984). Some more reddish chestnut in summer. The un- spots may persist hardly visible until late derparts are white, except the neck, which is October along the vertebral columns, on both brownish all around. There is buff or white sides. One seen in mid-September (1999) on the throat, ears and muzzle. The ears are showed spots only on the flanks. Young tipped dark brown or black, whitish inside, males show “buttons,” short bony pedicels and there is a black labial spot (Guthrie, 1971) from which the antlers will grow during sum- on each side of the chin, where the ancestors mer until September. The reddish brown of our deer had a white canine tooth protrud- summer pelage is comprised of wiry hairs. ed (canines are present in primitive deer). The The adult fur in winter is comprised mostly nose pad is glossy black. A white ring of hair of long, thick guard hairs. The gray is shed encircles each eye. from May 15 to June 15. Rarely one finds among white-tailed deer an albino or partially albino (piebald) pattern. Albinos are legally protected in Wisconsin. Dahlberg and Guettinger (1956) mentioned a melanistic deer in Vilas County, in 1948; another was seen in northwest Columbia County (Wozencraft, 1979). The antlers in mature adults may have as many as 12 tines, called points. Rarely a buck may have more, but usually fewer (in the more numerous younger bucks). A huge rack hanging in the mammal collection room here at this University has 13 points. It is marked Wisconsin on the wooden base. I observed a six point buck in velvet on Washington Island on July 1, 2003, one day after finding a newborn fawn. Antlers are shed in winter to conserve energy. Young- of-the-year males only have “buttons”, but Skull of Odocoileus virginianus. yearlings may have longer branched

TAXONOMIC ACCOUNTS / ORDER ARTIODACTYLA 441 “spikes”. The “rack” develops larger size in eastward of Wisconsin. Diminished in num- subsequent years (5-7 years old), but in aged bers greatly in pioneer times, the deer today bucks they grow to smaller size. Nutrition is found in every county, even in urban areas. and heredity effect how large a rack grows Because they swim well and cross the ice in (Smith, 1991). Prior to 1956, the average winter, they occur on most islands in Lake rack size was seven points. That has de- Superior and Lake Michigan. Sometimes they clined; because hunters have killed many are numerous and endanger rare plants in the large bucks. islands (Judziewicz, 2001). In 1999, near Ashippun, in southern Status. Once imperiled (Jackson, Wisconsin, Mike Peirick shot a 30-point buck. 1961; Schorger, 1953; Bersing, 1966; and There were 14 tines on one side, and 16 on others) by overhunting and habitat loss, the the other. The antlers were not typical, as management of white-tailed deer is a great branching was in clusters (five or six in one success story. Cory (1912) reviewed the cluster). The buck was not especially large, status in Wisconsin at the turn of that cen- but the rack was the fourth or fifth largest tury. Deer were more common in north- from Wisconsin. Considering the annual har- ern counties, but scarce in St. Croix, Dunn, vests in recent years (see below, Status), this Pepin, Eau Claire, Jackson, Monroe, Ju- unique buck was roughly a once-in-a-million neau, Sauk, Adams, Wood, Portage, Sha- opportunity. The cause of such multiple wano, and Waupaca counties. There were branching of antlers is unknown. only a few doubtful records in southeast The chromosomes are 2N= 70, FN= 70. Wisconsin, in Fond du Lac, Calumet, and Smith (1991) reviews other genetical charac- Manitowoc counties. Deer were rare in teristics of this species. There are two pairs Door County. The one-buck law was es- of inguinal mammae that superficially resem- tablished in 1915. By 1929, deer became ble an udder. Long and Smart (1976) com- common in Jackson, Monroe, Juneau, pared the os cordis, a small bone in the heart, Adams, perhaps in Calumet, and along the in whitetails from Wisconsin and Texas. They Wisconsin River into southwest Wisconsin. are larger in males and significantly larger in Deer occurred in Door County. But no deer Wisconsin. Atkeson and Marchinton (1982) were found at all south of the principal described the forehead glands. Metatarsal range in the northern counties. By 1938, glands are less than 25 mm long in the white- according to Walter Scott, there was little tail, longer in the mule deer. change, and deer were even more rare Total length varies up to 240 cm (8 ft) in along the southwest Wisconsin River Val- males, which weigh as much as 135 kg (300 ley. By 1954, the range (habitats) had pret- lbs), but seldom exceed 220 pounds. The ty much been restored, and deer were state- length is usually 140-200 cm. Females weigh wide in scattered occurrence. 20-40 percent less than males. Statewide, more than 200,000 deer are Dental Formula. I 0/3, C 0/1, P 3/3, M harvested each year, and 470,000 were har- 3/3 = 32. Usually there are no upper canines, vested in 1995. In addition, 40,000 were road or upper incisors, which are instead replaced kills. A similar number was taken in 1999. with a horny pad. The analogs are present in About 600,000 were harvested in 2000. The the lower jaw, and the canine has moved for- financial benefits to the state generated ward, compared to those in ancestral fossil through hunter’s licenses, lodging, gear, and deer, to become incisorform. traveling are incalculable but immense. In Geographic Distribution. This race of Wisconsin’s Deer Management Program (Bar- large, but small-toothed, whitetails ranged telt et al. 1994) these statistics are provided: from central Minnesota, all of Illinois, and “Assuming 50 pounds of meat per deer at $2

442 THE WILD MAMMALS OF WISCONSIN per pound of a Wisconsin harvest of 325,000 sequence of 5 severe winters in 8 years... deer equals $32.5 million in venison steaks, As recently as 1990 and 1991, populations sausage, and brats.” Add to this the recre- were above 400,000 (in record mild win- ational expenditures, and “the deer hunt to- ters beginning 1886-87). Populations de- day is worth at least $287 million.” Today clined in 1992 (due to antlerless harvests this may be double. and severe winter. Public opposition to lib- The geographic range and density of eral deer harvests in 1993 and 1994, re- whitetails no doubt would decrease eventual- sulted in an overwinter population again es- ly, if hunting were not allowed. The range, or timated at 400,000 during winter 1994-95. habitat, is all-important, and overpopulation The Wisconsin deer population now is larg- will destroy deer range. In Wisconsin, from er, but under constant management to gain 1949-1951, hunting pressure increased and the most from the ever-changing balance deteriorating range improved itself, reportedly of range and numbers. causing the state herd to increase. Since the Aside from the annual harvest there are 1940’s, the deer population in the north other benefits from deer. Deer save natural seems to be somewhat reduced. Among oth- habitats indirectly even for other species by er factors, increased tree growth and shade virtue of preserved hunting lands. Deer are reduced carrying capacity. Hard winters and food for wolves, coyotes, the rare Canada lynx deep snow impact deer populations by mor- and an occasional bobcat. Their carcasses are tality. Bad winters in succession or mild weath- carrion fed on by many birds and mammals. er patterns, such as the last three winters Often deer are nuisances, ruining or dam- through 1999, also affect deer numbers. In aging gardens, nurseries, orchards, and crops. many parts of Wisconsin human tolerance There is, then, an economic expense to many, determines the abundance. In the south where and a responsibility to society to pay the farm- small farms are the rule, deer and deer dam- er for such depredations ($15,000,000 damage to crops are not tolerated (i.e., they are age in 1993 alone) (see Bartelt et al., 1994). often poached). There is also the expense of car accidents McCaffery (1995) wrote: “The popula- ($92,000,000 annually in these years stud- tion fell from about 400,000 in 1964 to ied), not to mention injuries. Tree plantings fewer than 200,000 in 1972 following a and forest development are damaged by

Carrying capacity Central Wisconsin forest deer, 1962- Sketch of white-tailed deer stag. Leon L. Pray, in Cory. 1993. Current populations are too high. Barteldt et al.

TAXONOMIC ACCOUNTS / ORDER ARTIODACTYLA 443 browsing deer (Swift, 1948) and in nature sible epidemic of Chronic Wasting Disease is diverse wild plants are eaten (Vander Zouwen the problem of charitable use of venison, and Warnke, 1995). which some people fear to process, donate, Quotas for harvest are established on the accept, or eat. basis of deer numbers and carrying capacity Habitat. White-tailed deer have not (see Figure). That capacity is learned from changed their overall distribution in Wiscon- prolonged study. In the northern forests it is sin much, although they have dramatically about 15/square mile. At this density about increased. Favorable habitats include: forest 3.4 antlered bucks and the same number of edge, the mosaics of field and woodland (a antlerless deer may be harvested without ad- Wisconsin dairy farm heritage), preservation versely affecting the population (Bartelt et al., of public hunting lands, extensive marshes and 1994). In farmlands 100 deer/square mile is other wetlands, and several other kinds of not unusual, and some densities are even high- patchy habitats of land use (logging, small er. A typical quota of antlered or antlerless farms) (Gladfelter 1984). Where single crops deer is established for preservation of enough are the rule the yield is decreased. In winters does (which carry the next generation of deer). of severe cold and deep snow, deer struggle About 2/3 of antlerless deer are adult does. to find food and often starve or fall prey to The Department of Natural Resources enemies (see below) including free-ranging aims for a lower deer harvest than maximum domestic dogs. Deer also are found in park- because the populations at carrying capacity lands, even in cities, and especially in winter inflict extensive and irreparable damage in the in deep forests. River bottoms and brambles ecosystems. Other organisms as well as the are good places for them to hide. Corridors, whitetail itself may suffer population declines. ditches, and river bottoms are good places The goal for Wisconsin harvest is carefully for deer to move from one habitat to anoth- established by mathematical theory on the one er. Automobiles on highways kill so many deer hand and practical field studies on the other, that the highway may be considered adverse with the invaluable data of harvest numbers, habitat, and signs are regularly placed mark- to obtain a maximum sustainable harvest (see ing deer crossings. figures on winter severity and carrying capac- Areas dominated by aspen, oak, and jack ity). Studies are now in process and in plan- pine (or barrens habitat) can produce densi- ning to determine the effects of deer on the ties of more than 30 deer/ mi 2, and pole- total ecosystem (Bartelt et al., 1994; Vander sized red and white pine habitats, about 20 Zouwen and Warnke, 1995). deer/ mi 2. Other shaded and lowland habi- In recent years, urban deer populations tats support fewer than 15 deer/ mi 2 with have exploded in numbers. The Milwaukee extensive areas of pole-sized maples support- area receives much publicity because of dam- ing fewer than 10 deer/ mi 2. Canopy gaps age to gardens. Even the small city of Stevens in the original hemlock-hardwood forest Point (population 23,000) has been adverse- (1880’s) would have allowed in more sunlight ly affected. The occasional deer in the city than the shady northern hardwoods (mainly has been replaced in the past two or three sugar maple), that create low deer carrying years by small herds ranging into lawns and capacity today” (McCaffery, 1995). gardens. These deer, although making a great The whitetail has no conspicuous territory, nuisance of themselves, are often appreciat- but will defend a particular bed site from an- ed too. The state Department of Natural Re- other deer. In winter when the snow is deep, sources works with local communities to share the tree canopy reduces snow cover. Many the costs of managing these herds. The ven- deer tend to congregate in “yards” where the ison goes to charity. A side effect of the pos- snow is not so deep. Deer tend to lie on a

444 THE WILD MAMMALS OF WISCONSIN shallow depression, which can be seen in deep ern forests. Ironwood (Ostrya) and Cedar (Ju- grass or snow, raising their heads aloft now niperus) are also important browse plants. and then to look about. I have observed three Many Wisconsin deer eat corn in the does excavate snow forms, to the depth of fields and occasionally raid vegetable and flow- their bodies, using the front legs alternately er gardens. In some areas, 50-78 percent of in five or six rotary thrusts to the side. The the diet is from farmer’s crops. The quality of young fawn is never in a particular natal site venison is improved, so some farmers plant long, but lies alone, head down, hidden in corn for deer. ground vegetation flecked with sunshine, When deer yard up in the northern for- which matches the dappled coat. The fawn is ests to escape the problems of heavy snow, attended by its mother, 2-3 times daily, to the diet is focused on available woody vege- nurse and move it from one place to another. tation (Westover, 1971). In hemlock-hard- Foods. Deer spend most of the day, es- wood forest, on the Upper Peninsula of Mich- pecially at twilight and dusk, foraging for igan, a 13.7 square mile yard had an estimat- food. Forbs, grass shoots, and succulent ed carrying capacity of 14 deer per square leaves are eaten in spring and summer. Sum- mile. Only 10 lbs. of browse (over-dried sap- mer foods are documented by Dahlberg and lings up to six feet height) was available per Goettinger (1956) and McCaffery et al. acre. Ninety percent of it was stems [seed- (1974). Acorns, beech nuts, berries, forbs, lings and buds] of sugar, red and striped ma- and tree leaves are eaten in fall. In winter, ple Acer pennsylvanicum. Approximately leaves, cedars, grass, twigs and buds, even three pounds per day are essential amounts. woody browse are necessary. In the winter of 1968-69, the yarding popu- Winter starvation is not necessarily den- lation was estimated at 22 deer per square sity dependent (Vander Zouwen and Warnke, mile and the snow was deep, burying many 1995; McCaffery, personal corr.). When deer maple seedlings. At least 40 deer died that numbers are below carrying capacity starva- winter, and most autopsied had starved. In tion is seldom seen. The rate of reproduction 1969-70, a winter of moderate snowfall and may decline, with some pre-natal loss and a population of only 13 deer per square mile, mortality of fawns. On islands as well as in only one deer died (of unknown cause). Mor- the northern counties, overpopulation occa- tality was increased by the combination of high sionally impacts vegetation. population size and heavy snow cover. Vander Zouwen and Warnke (1995) list Reproduction. Breeding occurs from over 150 species of herbs potentially eaten October to January. Dahlberg and Guetting- by deer. They list numerous plants studied er (1956) review breeding information for showing that some are adversely affected by Wisconsin. Breeding peaks between Novem- deer, whereas other plants increase in number 10-29, when about 62 percent of the bers. Woody plants preferred (Swift, 1948) does are bred. A month later a second peak include white cedar, eastern hemlock, bass- occurs, during the next estrus. Fertilization wood, white pine, yellow birch, sugar and occurs in the uterine horn, and implantation red maples, aspen, and white (bur) oak, and about a month later. The embryos can mi- shrubby yews (Taxus) and hazel. Browse of grate from one horn to the other on occa- twigs may be as high as approximately 7 feet sion (Moore and White, 1971). Gestation ( = 2.1 m). lasts about 202 days (Hall, 1979), and var- In winter, yew (Taxus canadensis), bal- ies from 187-222 days (Verme and Ullrey, sam fir (Abies balsamea), hemlock (Tsuga 1984). At birth the newborn weighs about canadensis), and maple seedlings and sap- 3-4 kg. If nutrition for the mother is poor lings (Acer sp.) are important browse in north- the fetus will not grow well and may be re-

TAXONOMIC ACCOUNTS / ORDER ARTIODACTYLA 445 sorbed. Litter size varies from 1-3 fawns, and some of the populations. No doubt emigra- twins are typical. The newborn will double tion reduces density in one place, increasing its weight by two weeks of age. The spots it at another. on fawns are in four dorsal rows, and each Hunting, taking does, and other aspects spot is about one cm in diameter. There may of deer management are discussed by Smith be more than 600 spots on a fawn. The spots (1991). Vander Zouwen and Warnke (1995) are lost by the advent of winter molt. By 9- and Bartelt et al. (1994) discuss taking antler- 10 months the permanent teeth have erupt- less deer. ed. In a few weeks to two months, the fawn The wolf, coyote, black bear, and lynx, begins to graze on grasses and forbs. Wean- as well as the extirpated mountain lion, were ing occurs by 10 weeks of age (Smith 1991). natural predators of deer. Today, especially Verme (1965) studied diet on penned deer in deep snow or in southern states, the do- to determine adverse effects of poor nutri- mestic dog is a significant non-human killer tion on doe reproduction. This relation of of deer (Smith, 1991). Coyotes have become nutrition to physiology is reviewed by Ver- important predators of deer, responsible for me and Ullrey (1984). as high as 80 percent of fawn mortality in Females can breed at 6-7 months of age. some places. The return of the wolf and Can- Males breed in their second year. Nutrition ada lynx perhaps will affect the numbers of and photoperiod control the advent of annu- northwestern herds in Wisconsin. Predation al breeding. Life expectancy may exceed 20 culls out sickly animals and is generally of years, but few deer live beyond 10 years in benefit to the deer populations. the wild, and 2-3 years is an average life. Parasites include the deer louse Tricho- Late conception is adverse to fawns be- lipeurus and a blood sucking Solenoptes cause they have a month less time to devel- found rarely. Several ticks, including one op. Breeding in southern Wisconsin slightly that carries Lyme disease, are found on precedes that in the north. Birth of fawns is deer. Nose bots are common (Cephenemy- delayed in the north, so that more fawns are ia), but they cause the deer little harm. born before June 5 in central than in north- Worms such as Setaria, Wehrdikmansia, ern Wisconsin. I observed a doe with new- Gongylomema, Protostrongylus, and oth- born fawn on June 30, 2003, on Washing- ers are found in deer, and the moose brain- ton Island, and another newborn fawn in worm (Pneumostrongylus) occurs in the Stevens Point on May 1, 2003. The latitude meninges of deer. This parasite is a danger is similar, but the Island is cooler in spring. to the rare moose (see account of that spe- Mortality. The primary source of mortal- cies). Trematodes such as Fasciola hepati- ity in deer populations is legal gun and bow ca and the gigantic Fasciolodes magna are hunting. Other causes of mortality by humans found in the liver, and probably contracted include poaching, road kills, fence entangle- from feeding on aquatic vegetation (Hessel- ments, and forest fires. In winter the domes- ton and Hesselton, 1982). Wisconsin deer tic dog kills many deer. Deep snow and over- have been remarkably free of diseases and population lead to starvation, largely prevent- parasites compared to deer studied in south- ed by hunting. See section above on foods. ern states (Dan Trainer, UW-SP, personal Although managed harvests keep the corr.). Diseases that affect deer include epi- deer numbers below carrying capacity, some zootic hemorrhagic disease, arboviruses, populations do not require hunting regula- foot and mouth disease, occasionally rabies, tion. In dynamic environments, deer may and other viral diseases. Bacterial diseases keep below this level. Land predators and include anthrax, tuberculosis, and brucello- other mortality prevent overpopulation in sis, and it would seem pneumonia. Foreign

446 THE WILD MAMMALS OF WISCONSIN diseases, both bacterial and viral, may be- has been known about 30 years, there is no come a future problem from accidental or great problem. The Wisconsin deer herd cer- intentional transmission. tainly may be in great peril if the mad cow In the hunting season of 2001, three deer epidemic in England and Scotland is a mod- were killed from the Mt. Horeb area that test- el. Quarantine, slaughter, burning, burial, and ed positive for Chronic Wasting Disease the prohibition of caracasses for making food (CWD), and subsequently eight more were for livestock were accomplished there. Test- killed with 500 additional deer added to this ing in Wisconsin must be done state-wide, and Mt. Horeb sample. At this writing, more than why not utilize fresh road kills? Testing has 24 deer have been found positive in this area been carried out on 1,000 deer since 1999, (eastern Iowa and western Dane counties). from 70 of 130 management units state-wide, Probably brought into the state for deer and and no other positive cases have been found. elk game farms (imported from infected herds The attempts to gain financial support for this in several western states), CWD is a huge problem should be related to other possible threat to the entire deer population and all its outbreaks of disease (anthrax, bovine tuber- economic benefits for Wisconsin. Already the culosis, hoof and mouth disease, West Nile state has set aside $4,000,000 to develop new virus all are timely threats against security in strategies to prevent further importations and America), for the potential horrors of epidem- spread of the lethal, infectious proteins called ics endanger the entire human race. Another “prions,” which attack the central nervous recent disease found in Wisconsin, of antlered system. The Department of Natural Resourc- bucks, is called cranial abscessation syndrome. es is attempting to extirminate and test ap- Home range and density. The annual proximately 15,000 deer in the Mt. Horeb home range of the white-tailed deer averages area. Fortunately for humans, there seems no 59-520 ha (148-1,300 acres), and seasonal danger to them from infection, but the nega- use seldom exceeds 3.2 km (2 mi) diameter. tive impact on the state economy by loss of Home ranges are rather linear in shape, re- deer hunters looms as a disaster for people flecting the availability of patchy food resourc- as well as our native deer. Three immediate es. In homogeneous vegetation, the home problems must be solved: 1. Can the deer in range is circular (Smith 1991). the Mt. Horeb area be exterminated; and can Yearlings disperse more often and to far- killing these deer eliminate all the sickly indi- ther distances than other age class deer. Fawns viduals? 2. Has CWD already spread to other have the smallest home ranges. Winter home areas in the state? 3. Will the prion patho- range depends on snow depth; deep snow may gens persist in the environment and lead to confine deer to a wooded “yard.” Adult males future recurrences? At this time no ones wander irregularly, whereas the females and knows how Chronic Wasting disease is trans- fawns move more directly from one food source mitted (Manwell, 2002). [However, mad cow to another. Seasonal migrations of 15.6-23.2 disease is transmitted by nasal secretions and km (9.6-14.5 mi) to find food or shelter are deer often lick one another, and not only does common in northern populations of whitetails and fawns.] To complicate matters, the huge (Marchinton and Hirth, 1984). state herd must be managed by mortality, the Remarks on Behavior. White-tailed deer population cut by about 200,000 deer annu- live in one of two basic social groups. The ally and harvested by hunters, who now may Matriarchal group includes a mature doe, her be afraid to eat the venison. Some say a great daughter(s) and fawns. When fawns are young, tragedy is impending, such as the mad cow the mother and fawns comprise the social epidemic in Great Britain, also caused by pri- group. In two years, I observed two mothers ons. Others say that in Colorado, where CWD that usually grazed together with their fawns

TAXONOMIC ACCOUNTS / ORDER ARTIODACTYLA 447 as social groups. The males usually live sepa- Severinghaus (1949) (see Hoffmeister, rately in loose, fraternal groups including adult 1989) provides different (Illinois) catgegories and most yearling males. Sometimes Mixed- as follows: sex groups are found, but usually males seek Fawns: Interior deciduous incisors (I 0/ females only in the season of rut. Males are 1), less than 4 months. Replaced with per- highly territorial when the object defended is manent incisors, 5-6 months. Other incisors a potential mate. In the larger groups there replaced by 12 months. Second molar erupt- are dominance hierarchies that affect the being at 7-9 months. havior of the group members (Smith 1991, Yearlings:Third molar erupts at 12-16 Forand and Marchinton 1989). They mini- months. Permanent premolars appear 18 mize conflict and reduce expenditure of ener- months later. Any dP’s retained are badly gy in the winter. The communication is by worn. Thus, the molars erupt somewhat ear- stereotyped postures and other signals. Males lier compared to the above example. in rut often engage in fighting, more or less Adults: 2 1/2 year-olds have lingual strenuously in my observation, to obtain dom- crests of molars sharp, with enamel well above inance and to mate a particular doe. the brown dentine line. Wear on M /3 is slight. “Buck rubs” and scraping (Sawyer et al. 3 1/2 year-olds have a posterior cusp of M / 1982) are visual and olfactory signposts of the 3 worn flat. 4 1/2 year-olds. M /1 showing older males (Kile and Marchinton, 1977, Ben- much wear; P /1 showing little. 5 1/2 year- ner and Bowyer 1988). During rut the fore- olds. Molars worn and p /1 shows some wear. head glands, salivary glands, and preorbital 6 ½-year olds show ground surfaces and P /1 glands all contribute odors to the rubbed shows moderate wear. Older deer show “cup- branches (Atkeson, 1982). A buck “rub” is a ping” of cheek teeth. stem or sapling, often broken, pulled down by Fetuses may be aged by crown-rump a buck, and scented with the head gland. A length, or when longer, a straight-line distance buck “scrape” is where a sapling is pulled down, between the forehead and rump. From Dahl- often broken, stripped of bark, and around it berg and Guettinger (1956), based on earlier the deer paws a depression. Usually he uri- work by Armstrong. Cheatum, and Morton: nates in the “scrape” (Marchinton and Hirth, Lengths in millimeters and ages in days were 1984). Bawling and bleating to call the moth- 20/40, 30/45, 40/50, 50/54, 60/58, 70/ er are mentioned by Smith (1991) and Atke- 61, 80/65, 90/68, 100/72, 110/75, 120/ son et al. (1988), and I have heard fawns bleat. 78, 130/81, 140/83, 150/86, 160/88, Remarks on Aging. Deer are aged by 170/91, 180/94, 190/97, 200/100, 210/ tooth eruption and wear, which vary some- 103, 220/106, 230/109, 240/112, 250/ what from place to place. In any locality a 115, 260/118, 270/121, 280/124, 290/ series of jaws properly aged will, when used 127, 300/130, 310/133, 320/136, 330/ as a standard, allow the local development 139, 340/142, 350/145, 360/148, 370/ and age parameters of the deer population to 151, 380/154, 390/157, 400/161, 410/ be studied. Usually by four weeks there are 165, 420/170, 430/174, 440/179, 450/ four incisorform teeth, two on each side (I 0/ 185, 460/192, 470/203, 480/214. If plot- 3, C 0/1). Two upper premolar teeth also ted, these increments vary from 3-5 mm each will have erupted. All of these are deciduous. 10 days until 91 mm length is attained, then By seven months the first permanent molar the growth is steady (3 mm/10 days) until 154 has erupted. By 13 months the second ap- mm is attained, then growth accelerates from pears, followed shortly by the third. The teeth 4 to 11 mm/10 days until 214 mm length is are fully erupted about 19 months of age (Hes- attained. Obviously such estimates are affect- selton and Hesselton, 1982). ed by starvation of the mother, multiple births

448 THE WILD MAMMALS OF WISCONSIN or single offspring, food scarcity and avail- display from a doe to lead predators away from ability, and disease. concealed fawns (Smythe, 1977); and cohe- Remarks on color patterns: The spots on sion effect or group alarm signals, presumably fawns are obviously adaptive in concealment to allow deer to escape in groups. coloration, and fit well with the hiding behav- Remarks on threats in young deer. I ior of fawns. Explanation of the white flag’s present some information on the development evolution is controversial. Such an aposemat- of threat and fighting of deer having velvety ic signal may distract attacking predators. Hirth spikes or near the time of parturition. Little and McCullough (1977) believed the alarm sig- has been written about the onset of antler nal also warns other deer of the predator. Oth- growth and associated behavior in whitetails, ers have speculated that the flag is a notice to except their use by adult males with mature the predator that the deer sensed it and there- antlers. I examined at close hand (by 17 May by has informed the predator of this, causing 1999) a deer with short spikes in velvet, one a the predator to look elsewhere for food. When gray stub approximately 100 mm long, the a neighbor’s husky dog suddenly startled up a other already branched and showing signs of matriarchal group of seven deer in my back- further branching at the tips of the tines, more yard, there were flags going up everywhere. than double the length. This deer had a tiny Two late fawns struck out one way, while two “mane” from the crown of the head extending others and the adults went off another. The onto the neck, and a smaller analogous “mane” flashing of flags reminded me of fireflies that on the mid-line of the chest. On 15 May 2002, flash synchronously to inform the nearest I observed a pregnant doe and two males with potential mate where to be found, with a stubs approximately 2 inches (51 mm) in minimum of confusion. Not-in-synchrony flash- length. These two were chasing each other and ing should work the opposite, i.e., cause con- the doe, leaping in and out of the waters of fusion to the flashers. As for evolution of elon- McDill Pond, occasionally drinking, and often gate white flags, one should note that the un- fighting with forefeet and “stub” antlers. One derside of the primordial tail was already white, wading in the water stood on its hind legs re- and the white could be concealed or present- peatedly and flailed at the other standing above ed. It is a simple evolutionary change to elon- him on the bank, actually striking it once or gate a bicolored tail into a conspicuous flag. twice with a hoof of the foreleg. The next day There are five explanations given for flagging: it was seen walking with [probably] the same aposematic startling and confusing of preda- doe, which gave birth a few days later. The tors; alarm signal to other whitetails; invitation male was seen no more. In June 1999, I ob- to predators saying, in effect, you have been served a few-days-old fawn making head threats seen and I have already got away; distraction to a mallard hen (defending her young on a path), and it chased them into the water. Geographic Variation. There is one race in Wisconsin. Additional Life History. There are countless works on the whitetail, both popular and technical. An old one in Wisconsin is by Dahl- berg and Guettinger (1956, The White-tailed Deer in Wisconsin). An important monograph on the Cervidae is Kellogg (1965) in Taylor’s The Deer of North America. Dahlberg and Guettinger (1956) reviewed the biology of the Whitetails flashing in synchrony. Artist Dan Metz. whitetail in Wisconsin up to 1956. Halls

TAXONOMIC ACCOUNTS / ORDER ARTIODACTYLA 449 (1984) is a classical work on this deer, and vomer incompletely dividing the narial cham- Wemmer (1987) treats the family Cervidae. ber, and a wide lacrimal or antorbital fossa, Specimens examined. Total, 19. Bay- the metatarsal gland of Odocoileus is absent field, Door, Dunn, Langlade, Manitowoc, in Alces, and much of the hair is buoyant, and Marinette, Marquette, Portage, Price, Wau- brown or black dorsally. The front hooves are paca, Waushara, Wood counties. enormous. Usually the moose is solitary (espe- Michigan: Little Summer Island 1. cially in winter). It is often diurnal.

Alces alces (Linnaeus) Moose

Alces alces andersoni Peterson

1950. Alces americanus andersoni Peterson. Oc- cas. Papers Royal Ontario Mus. Zool., 9: 1. Type from sect. 27, Sprucewood Forest, 15 mi. E Brandon, Manitoba. Palmate moose antler. 1952. Alces alces andersoni: Peterson. Contr. Roy- al Ontario Mus. Zool. and Paleontol., 34: 24 1983. Alces alces americana (Clinton): Baker. Mich- Genus Alces Gray, 1821 igan Mammals, p. 599. Moose The name Alces means elk, which is what The moose is a large mammal, the largest Europeans usually call this mammal. Some at- deer in North America, slightly larger than tribute the word to the Greek (alke), which means the elk. It has long forelegs, short tail, broad strong. Moose is derived from an Algonquian muzzle overhanging the lower jaw, a dewlap Indian word moos, or musee, a browser of twigs. present on the throat, and antlers in males Description. See account of the genus. are heavy, palmate, i.e. with relatively short American moose have a karyotype of 2N=70 points. The skull has premaxillaries elongated, (Coady, 1982). There are two pairs of mammae. There is no metatarsal gland. The color ranges from tan to nearly black. In winter it is grayer. Molt takes place in spring, when short and dark-tipped guard hairs grow out, reaching in summer 20 cm or more in length. Young are red-brown and lack spots. The head and body length is 240-310 cm (approximatelly 2.5-3 yds). The weights, especially in Alaskan moose, may reach 825 kg (1.833 lbs). Coady (1982) gives weights as 418 kg for males (929 lbs) and 414 kg (920 lbs) for females. Peterson (1974) summarized data on weights for both sexes. .See comments in Re- production below. The length of the skull may exceed 600 mm, smaller in females, and the Moose cow. Artist Caton. Deer in America. maxillary tooth-row as long as 150 mm.

450 THE WILD MAMMALS OF WISCONSIN Dental Formula. I 0/3, C 0/1, P 3/3, M moose meningeal worm seen commonly in 3/3 = 32. white-tailed deer, 2. Presence of suitable hab- Geographic Range. Usually found in itat, marshy wetlands with willows and other northern counties of Wisconsin and Upper riparian shrubs for browse. 3. Remote habi- Michigan, but occasionally the moose wan- tat away from roadways. 4. Absence of ders southward as far as Portage, Green Lake, poaching. Another danger may be the new- and Sauk counties. Unknown from the Door ly discovered Chronic Wasting Disease, Peninsula. Schorger (1957) mapped out its which infects whitetails and elk. The moose historic range. After its extinction in Wiscon- has little contact with whitetails, but it be- sin in 1921 (and it was extirpated much ear- longs to the same family Cervidae. I believe lier from this state than indicated by that late the moose can return to Wisconsin so long Minnesota record), for perhaps 50 years there as it is given protection. were no records of moose in Wisconsin. The Habitats. See Distribution and Status moose reinvaded the northern counties, prob- above. Boreal forests (where snow covers the ably from Minnesota, and has almost reached land in winter) and extensive wetlands, where the limits of earlier breeding range. willows and aspens provide twigs for browse. In 1993, a moose was killed by an auto- Aquatic vegetation is also eaten, even from the mobile in Stevens Point. It had wandered bottom of a stream. The twigs are preferably about in the nearby Dewey Marsh most of in early tender development, but in winter hard- the summer, where it was seen several times er branches are eaten. Browse includes shrubs that year and the year earlier. The specimen of many species, willows, aspen, balsam fir, was the first in central Wisconsin since the paper birch, and much muskeg vegetation in- turn of the century (see Schorger, 1957). A cluding mosses. In winter, moose may move recent questionable observation in 2003, was to riparian woodlands from the marshes. Log- from Calumet County. ging and fires are detrimental to moose, al- Status. The Minnesota moose population though they readily reinvade seral habitats. has increased since protection was initiated in Their thick fur is buoyant, their preferred hab- 1922. Recent movements show invasion into itats wetlands. Moose swim well and may fend southern Minnesota. Moose in Wisconsin, where off wolves after entering water. it is protected, are becoming more common and Foods. See habitats above. are breeding here. There have been some cows Reproduction. Females in the season of with calves reported near Rhinelander in recent rut (September to October) moan, and the years. Cows were seen with calves in Ashland males croak and push one another around with and Forest counties (Wydeven and Wiedenhoe- their immense racks. Yearling males are capa- ft, 2004 report). Wydeven (1993) compiled ble of breeding but the older bulls may not al- records of several other sightings in Wisconsin, low it (Peterson, 1974). If the female has not which are included here as follows: Douglas, bred, one estrus period follows another at in- Bayfield, Ashland, Vilas, Forest, Washburn, Saw- tervals of 18-30 days, usually around 21 days. yer, Portage, Shawano, and Oneida counties. One bull may follow a cow for a week or more He estimates that there are presently 20-30 and then turn his attention to another. The moose in the north. The Portage County moose calves born usually result from one estrus, with was a bull weighing about 800 lbs. It was ap- rare births resulting from breeding at earlier or proximately 122 km and 170 km southward of later times. Estrus lasts about a day. Late births Wydeven’s nearest records (Fifefield, Price Co., from late estrus result in calves having restrict- and Woodboro, Oneida Co.). ed development. Most calves result from ma- The return of the moose depends on ture breeding animals 5-11 years of age. Ges- several factors: 1. Little adverse effect of the tation is approximately 243 days, and calving

TAXONOMIC ACCOUNTS / ORDER ARTIODACTYLA 451 takes place in late May or early June. Calves to moose it is often a lethal parasite. Although follow their mother after 2-3 weeks, and wean- unknown in moose, chronic wasting disease ing takes place by 5 months. Both sexes can of whitetails may infect moose. Flukes, tape- mate in their second year. Seldom are twins worms, and ticks are other parasites. or triplets produced by yearling cows. Usually Home Range and Density. Home range there is a single calf, which weighs about 11.2 may be variable for one moose from year to kg at birth, born with fur (Verme, 1970). Twins year. It seldom exceeds 5-10 km2, and is even are not uncommon. High densities of moose smaller in winter. For 25 days a Minnesota result in a lower reproductive performance pre- moose stayed in a 2.4 km2 balsam fir stand sumably because females have fewer resourc- (Van Ballenbergh and Peek, 1971). Moose es. By October a young moose may weigh 180 wander in summer sometimes haphazardly, kg. In captivity a moose probably might live and for great distances. Most movements are well over 20 years, but in the wild it has been made in search of food. In Wisconsin, obvi- documented only 8-10 years (Crandall, 1964). ously, there is little information about density Data on growth was obtained by J. or home range. Moose may aggregate in M.Peek, R. H. Denniston, G. Markgren (for groups in fall and winter, but usually the ani- Swedish moose) and J. M. Heyland, and were mal is solitary unless the animal is a mother. summarized by R. L. Peterson (1974). Additional Natural History. Coady (1982) Data on moose calves from 2-91 days is an excellent reference source for the moose. show growth in total length to 162 cm, and Geographic variation. None is observed nearly 60 kg in weight. Adult male and female in Wisconsin. moose obtained from 1962-1966 in Quebec Specimen examined. Total 1. Portage harvests varied by age class as follows: Co. Stevens Point 1. Other recent records (Wydeven 1993): Age N Mean Weight (kg) Douglas, Bayfield, Ashland, Vilas, Forest, males females Sawyer and Oneida counties. 6 months 19 166 kg 26 50.7 1 year 51 283.6 34 280.7 II 27 361.7 16 332.3 III 49 386.3 22 348.1 IV 50 456.2 35 361.7 V 39 496.1 34 389.3 VI 32 487.9 34 368.9 VII 48 491.2 29 364.2 VIII 19 511.6 17 385.6 IX 36 475.8 7 332.1

These data suggest moose grow until the fifth year, and possibly decline in weight somewhat in the ninth year. Mortality. The chief enemy is humankind by both hunting and automobiles. Wolves kill moose (Mech 1966). Bears and the Canada lynx may occasionally take calves. Occasion- ally a rutting male may kill its rival. The meningeal worm Paralaphostrongylus tenuis Map showing geographic range of Alces alces in North is often contracted from white-tailed deer, and America.

452 THE WILD MAMMALS OF WISCONSIN GLOSSARY OF Baculum- A bone found in or extending into MAMMALOGICAL TERMS the glans penis, also called os penis. Basioccipital — The bone of the occiput ly- Acrocentric — A chromosome having the cen- ing below the foramen magnum. tromere toward one end. Basisphenoid — A bone lying anterior to the Adult — Mature age class. Identified by sexu- basioccipital. al development, indicated by tooth wear, Beam — The main branch of an antler. fusion of bone sutures, etc. Bergmann’s Rule — Mammals tend to large Age class — One stage from birth to death, body mass in cold habitats. as juvenile, young, subadult, adult, old Bicornuate — Having two “horns” or pro- adult, or class I and class II shrews. cesses as a type of uterus. Each oviduct Albino — Whitefurred. True albinos have pink is the cornu, or “horn”. eyes. Biomass — The organismal mass, including Allen’s Rule — In hot, dry habitats append- all individuals in a given area. Usually cal- ages (ears) are larger; in cold habitats they culated as dry weight per given area. are smaller. Biome — An ecological area defined by the Alpha male or female — Dominant male or dominance of certain important plants female, usually the sole breeder in its or animals, Tall Grass Prairie or Spruce group. — moose biomes. Altricial — An ornithological term used for Birth rate — Number or offspring produced helpless young. in a given time by a given population (of- Alveolus — Tooth socket. ten a theoretical 1,000). This is the crude Alveolar length — Longest dimension of a rate; the specific rate is that of females tooth socket, measured at the rim. of given age. Angular — Most ventral of three posterior pro- Bradycardia — A significantly slowed heart cesses of the dentary (lower jaw). rate. Annulus, annuli pleural — Annual deposit of Breadths — See widths. cementum or calcium in a tooth. Browse — leaves, buds, and twigs of woody Antler — Paired bony extensions of frontal vegetation used as food by deer. bones that grow covered with skin and Bulla — A bony capsule: usually called audi- hair (“velvet”), usually branched and seen tory, otic or tympanic, but also applies only in the Cervidae. to the mastoid. Antorbital canal — See infraorbital canal. Bunodont — A tooth, usually a molar, with Articular — The middle of three posterior softly-rounded cusps (on upper teeth) or processes of the dentary. Articulates with cuspids (lower). squamosal. In carnivores functions as a Calcar — A stiff cartilage extending along the pin-hinge and is called condyloid process. margin of the bat’s uropatagium from Arvicoline — Formerly called Microtine (see each hind foot. below). Can trap — A can set into the earth flush Astragalus — A large bone of the ankle next with the surface, used to capture shrews. to the heel bone (calcaneus). Also called a pit trap. Auditory bulla — A thin bony capsule overly- Canine — A fang. Elongated first tooth in the ing the middle ear comprised of the ec- maxillary bone used usually for stabbing totympanic ring and usually an endotym- or slicing prey. panic inward extension. Cannon bone — A pair of metapodial bones Auditory ossicles — Bones of middle ear: fused longitudinally, seen in the order Ar- Malleus, incus and (innermost) stapes. tiodactyla.

GLOSSARY OF MAMMALOGICAL TERMS 453 Carnassials — The major carnivoran shear- Delayed implantation — Fertilized blastocysts ing teeth, each upper fourth premolar do not implant to develop until favorable and lower first molar. conditions. Carnivore — A member of the order Carnivora Density — Number of individuals of a species that includes dogs, cats, bears, and allies. present in a given area, e.g., 1/ha. Cheek pouch — An invagination or evagina- Dental formula — Numbers of teeth on ei- tion of the cheek for food transport. Ex- ther side, categorized as incisors, canines, ternal cheek pouches open to the out- premolars and molars for upper and low- side and are fur lined. er jaws, doubled for total. For Man, DF I Chorio-allantoic placenta — A “true” placen- 2/2, C 1/1, P 2/2, M 3/3 = 32. ta, which is found in Eutherian mammals, Dentary — The ramus of the mandible, this advanced beyond the marsupial’s yolk sac bone bears lower teeth, and is one side placenta. of the lower jaw. Cloaca — A commom opening for emission Dentine — Hard inner substance of the tooth of feces, excretory wastes, sperm, eggs comprised of heterogenous prisms of cal- (or newborn young). cium-phosphate. Coronoid — The uppermost of three posteri- Dew hooves — Paired vestigial hooves, per- or processes of the dentary bone. sisting above the inner functional hooves. Corpus luteum — A gland formed from the An analogous claw in some mammals is egg follicle in the ovary that produces a “dew claw”. progesterone following ovulation. One Diaphragm — Muscular respiratory organ ly- corpus luteum = one ovulation in corpo- ing transversely across the trunk, located ra lutea counts. immediately below the lungs and rib cage. Cranium — The bony housing of the brain Diastema — Gap in the tooth row, usually often called braincase, and often used as between cropping and chewing teeth. a synonym of skull. Digitigrade — Adapted to run on the toes. Crepuscular — Active in the evening, at twi- Dilambodont — Wshaped crest in primitive light. molars; also refers to such molars. Cretaceous Period — Time about 65 mya, i. Diphyodont — Two sets of teeth, deciduous e., 65 million years B.P., preceding the and permanent. Paleocene Epoch. Diploid (= 2N) — total number of chromo- Cricetine — Popular term for most New World somes in a body cell. Gametes are hap- non-arvicoline mice, but technically ap- loid, i. e., N. plies to Old World Cricetus and its allies. Dispersal — A straightline distance between Crista-- Crest of a tooth. place of birth and place of breeding. Cursorial — Adapted to running. Dispersion — Immigration into new habitats, Crown — Uppermost, usually referring to the extension of range. Some have used this enamel covered tooth exposed above the term for dispersal. gum. Diurnal- Active during the day. Cusp — A sharp or softly rounded upward Dormancy — A deep seasonal or regular sleep projection of the crown. characterized by only a slight drop in Death rate — Numbers of deaths per popula- body temperature. tion (often given as theoretical 1,000) in Dorsum — The back, the uppermost side. a given time. Duplex — Uterus divided into two separate tubes. Deciduous — Seasonally or developmentally Echolocation — Orienting locomotion by re- shed, as milk teeth or antlers. Trees that ception of echoes. Sonar of marine mam- shed leaves. mals is similar.

454 THE WILD MAMMALS OF WISCONSIN Ecosystem — An organism and its surround- Family — A taxon higher than genus and lowing living organisms and nonliving ele- er than order. The family name has the ments interacting independently of oth- suffix -idae. Note also subfamily –inae. er such systems, with the source of all Faunal Area — A realm or smaller ecological energy being chiefly the sun. area known by certain mammals being Ecotone — Area where two biomes or com- present or absent. munities meet and merge. Fenestra — A window; fenestrate means per- Ectotympanic ring — In the primitive mam- forated. mals the ear ossicles are protected by a Fertilization — Penetration of an egg by a circular ectotympanic bone. sperm. Edge — See text under Environment and Ecol- Fetus — Growth state developed before birth ogy. Interface of different habitats, e.g., but beyond embryonic, i.e., resembling forest and prairie. adult features. Emigration — Movement of individuals out Foramen (Foramina plural) — A small hole in of a population. a bone. Enamel — Exceptionally hard outer substance Foramen magnum — A moderately large comprised of calciumphosphate prisms opening in the occiput for the spinal cord. arranged perpendicular to the surface of Form — A cryptic depression made in the the tooth. snow, ground or grass in which mam- Endogone — An odiferous fungus on which mals rest and hide. some mammals feed. Fossorial — Adapted to digging, as a mole. Epiphysis — A bone developing separated Fractal — A geometric pattern based on a from another bone by cartilage; eventu- fractional exponent and an incipient cha- ally by ossification of the cartilage, the otic phenomenon describing complex epiphysis fuses to the other bone. scaling and irregularity in time, body EQ — Encephalization quotient, a ratio of log structure, or environment. Any tenden- brain mass/ log body weight, compared cy toward remarkable intricacy by scal- with normal points plotted on or along ing may be called “fractal.” the linear curve for mammals. Frontal — A bone in the skull roof above the Estrus — ”Heat”, i.e., sexual receptivity of orbit, housing the anterior part of the brain. female mammals. Fundamental Number (= FN) — The number Evolution — Changeover in the genes of a of biarmed and acrocentric elements of population usually observed over great autosomal arms in the didploid set of expanses of time. New genes eventually chromosomes, excluding the sex chro- may be selected for adaptive functions mosomes. and characters. See Speciation below. Genus — A taxonomic category above the External measurements — Also called stan- rank of species and below the family. The dard measurements, these are total, tail, generic name is capitalized, and under- hind foot, and ear lengths, in that order. lined or printed in italics. Extinction — the absolute eradication every- GestationThe period between fertilization and where of a taxon. birth. Extirpated — Eradicated, i.e., a kind of or- Gloger’s Rule — Mammals are pale in dry ganism entirely removed locally or re- habitats. gionally. Guard hair — Elongate, coarse hair project- Eye lens — The crystalline lens in the eye of- ing beyond the underfur. ten used for determining age (lens Habitat — The organism’s dwelling place, its weight). home and surroundings.

GLOSSARY OF MAMMALOGICAL TERMS 455 Haplodont — The primitive reptilian form of Insectivorous — Adapted to feed on insects. tooth, single cusped. Interfemoral membrane — A patagium be- Hallux — The first toe. tween the femurs, as in bats. Also called Herbivore- An animal that usually eats plants. uropatagium. Heterodont — Teeth which differ in form and Intergrade — Hybrid organism intermediate function. in characters of two adjoining subspecies. Hibernaculum — A place where an animal Interorbital — See widths. hibernates, a cave or den. Juvenile — An animal in the earliest phase of Hibernation — A sleep characterized by un- postnatal growth, usually characterized controlled temperature regulation, so that by milk teeth in mammals. Juvenal is an within limits the body temperature var- adjective describing such a juvenile. ies with ambient temperature. Karyotype — The chromosome picture for Holocene Epoch — See Recent Epoch. an individual showing all the pairs of chro- Home range — The area within which an mosomes. animal usually wanders. In practice, a Labial — near the lip or cheek. mapped area generally bounded by pe- Lactation — The process of producing milk. ripheral sites of captures and recaptures See Mammae. of a marked individual. Lengths — The Iongest dimensions. Total: Horn — An extension of the frontal bone Length from tip of fleshy part of tail to overlain with a smooth keratinous sheath. nose. Length of tail extends along tail Hybrid — Offspring from mating of two taxa from where it leaves the back to its fleshy such as species or occasionally genera. termination. Length of hind foot extends See intergrade. from heel (which is an ankle bone) to tip Immigration — Invasion into a new area. of longest claw. Ear length is measured Implantation — The attachment of the embryo from the notch to the distal tip of the (with its surrounding trophoblast cells) to pinna. Greatest length of cranium ex- the uterine wall for placental prenatal care. tends from the farthest posterior exten- Incisive foramen — One of paired palatal sion of the supraoccipital or occiput (usu- openings between the premaxillary and ally the condyles) to the anteriormost maxillary bones, serving as a taste area extension of the rostrum (nasals, premax- (organ of Jacobson) or as a canal for illaries, or incisors). Length of nasals is nerve and blood vessels. measured anteroposteriorly to the far- Incisor — Anterior teeth usually modified for thest extensions. The maxillary toothrow nipping, grooming the fur, and for gnaw- includes all the teeth except the incisors, ing in some herbivores. An incisor is ei- measured along the alveoli in the upper ther found in the premaxillary or is an jaw. The length of the palate includes the analog in the dentary. premaxillary but not the incisors, and ex- Infraorbital canal — A passage of the skull tends to the nearest site of the border of leading from the orbit and opening on the pterygoids. Occipitonasal length the rostrum. Usually contains a nerve and ranges from the anteriormost part of the blood vessels; muscles traverse this pas- premaxillaries to the occipital condyles. sage in many rodents. Also called antor- Lens weight — The weight of an oven or in- bital canal. cubator dried lens extracted from the Inguinal — Found on the posteriormost part eyeball, used for ascertaining age. of the venter, between the thighs. Lifezone — A temperature belt in North Insectivore — A member of the order Insec- America, sometimes identifiable on tivora, as a shrew or mole. mountains. In the American West from

456 THE WILD MAMMALS OF WISCONSIN uppermost Arcticalpine to Hudsonian, Metatarsal gland- A gland on the hind foot Canadian, Transition, Upper Sonoran, below the ankle, as in the Cervidae. Lower Sonoran. The concept was devel- Microtine — Tooth showing occlusal prisms oped by C.Hart Merriam and loops, as in Micrtotus. Lodge — A house made by beavers of mud Migration — A seasonal movement of an or- and sticks. ganism involving a round trip. Lord’s Rule — In related mammals litter size is Milk tooth — See Deciduous tooth. larger at higher latitudes. It has exceptions. Mist net — A net woven of fine black thread Lophodont — Tooth type with transverse crests. used to capture bats. Mammae — The teats, organs of lactation. Molar — A posterior permanent cheek tooth The nipple projects from the rounded that is never replaced or never replaces part sometimes called the mamma. another. “Mammary glands” are microscopic, sub- Molt — The change in fur, new replacing the dermal, and produce milk. old. A seasonal change in body hair Manus — The hand. Mortality — Death. Usually given as a ratio Marsupial bones — A pair of bones that ar- of killed individuals per population (of- ticulate with the pubic bones, only seem ten a theoretical 1,000). in the orders Monotremata and Marsu- Nasal — The anteriormost bone of the skull pialia. Also called epipubic bones. roof, overlying the olfactory chamber. Mass — In practice, body weight, in grams or Nematode — A roundworm, often an inter- other measures of weight. See Biomass. nal parasite of mammals. Masseter — Muscle that closes the jaws, with Neopallium — Wrinkled cerebral cortex seen origin mainly on the zygomatic arch and in many mammals and attributed to this insertion on the dentary. entire order. Mastoid Bulla — A portion of the socalled Niche — The animal’s complete biological “temporal complex” bulging through the occupation or “profession.” Also called skull wall lateral to the occiput. the role. Maxillary — A bone of the skull that partially Nocturnal — Active usually at night. covers the anterior part of the brain and Occiput — The posterior part of the skull nasal chamber, comprises part of the comprised of four bones surrounding the hard palate and houses the roots of all foramen magnum the upper teeth excepting incisors. Occipital condyles — Two condylar process- Maxillary toothrow — The length from the es of the exoccipitals, one on each side anteriormost part of the canine alveolus of the foramen magnum for articulation to the posteriormost part of the alveolar with the vertebral column. rim of the last molar. Ochraceous — Orange tan color of pelage. Melanistic — The fur is black owing to the Omnivorous — Adapted to feeding on both black pigment melanin. plants and animals. Metabolism — The chemical processes of the Orbit — The recess in the skull for each eye. body. There are Total (all), standard (what Order — A taxonomic category above the can be practically measured), basal (the family level, below the class. theoretical minimum, usually calculated) Os clitoridis — A small bone in the clitoris and relative (in relation to a unit of mass, homologous to the baculum. such as g). Os penis — See baculum. Metacentric — A chromosome arrangement Ovary — Organ in the abdominal cavity that where the centrosome is near the mid- matures and emits eggs, estrogen and dle, not at either end. progesterone.

GLOSSARY OF MAMMALOGICAL TERMS 457 Ovulation — The emission of an egg from meant to closely follow birth, e.g., post- the ovary into the uterus. partum estrus. Palate — The hard, bony shelf separating the Precocial — Born or hatched able to see and nasal chamber from the pharynx, com- move about. prised of premaxillaries, maxillaries, pa- Predator — An animal that kills and eats prey. latines, and pterygoids. Prehensile — Adapted to grasp, such as the Palmate — Describes antlers where the pro- opossum’s tail. jections are short and broadly connected Premaxillaries — Paired bones anteriormost in instead of long and spike like. Resembles the upper jaw, housing the upper incisors. the palm and fingers of the human hand. Premolar — Tooth used for maceration, weak Parapatric — The geographic range of one shearing or crushing, found between ca- mammal adjoins that of a closely related nine and molars. species, without overlap. See sympatry. Preputial glands — Invaginated skin glands Parietals — Paired dorsal roofing bones of near the penis. the skull posterior to the frontals and Primitive Number — Number of teeth in an- anterior to the occiput. cestral placentals, usually 44 (I 3/3, C Parturition — Birth. 1/1, P 4/4, M 3/3) X 2. Patagium — A membrane developed for glid- Quill — A specialized detachable hair stiff and ing flight; in bats comprising a complex pointed, for defense, in porcupines. of flight membranes. Rassenkreis — An overlap in a ring of races Pectoral — On the chest. where the overlapping (i.e., sympatric) Pelage — Fur, the hairy coat. populations cannot hybridize. This is the Phalanx (phalanges plural) — A bone of a toe commonest word that is used for this or finger. phenomenon. Pheromone — An external odor, a releaser or Reabsorption — Breaking down and removal primer initiating some particular behavior. of an embryo by the mother’s body. Philopatry — Group wherein young remain Recent Epoch — See Holocene Epoch. in the natal area, or with a parent. Present epoch after Wisconsin stage (sta- Pinna of ear — Paraboloid, fleshy projection of de) of the Pleistocene. outer ear. Sometimes called the ear conch. Reentrant angle — The wall of a molar tooth Placenta — Organ attached to uterine wall inclined inward as a sharply angled corner. for gas transport and nourishment of Retractible — Capable of being withdrawn embryo in mammals. from view, cat claws. Plantar tubercle — Protruberance on foot sole, Rhinarium — Nose pad. often mentioned for field mice. Root — Portion of a tooth set into the jaw bone. Plantigrade — Adapted to walking on the Rooted — Upon maturity the root closes off its plantar surface (sole) of the hind foot. blood vessels and permits no further growth. Pleistocene — Epoch of geological time prior Rootless — Evergrowing tooth. Root open to Recent, characterized by glacial and permitting blood vessels to facilitate con- interglacial stages. tinuous growth. Postmandibular foramen — In shrews, an Rostrum — Portion of skull extending forward opening on the dentary near the articu- of zygomatic arch. lar condyle. Rut — Breeding season of Artiodactyls, when Postorbital process — Pointed projections of males court females. the frontal or jugal bone. Sagittal crest — Longitudinal medial promi- Postpartum — Life stage, state or process nence of the skull for attachment of tem- following the giving of birth, usually poralis muscles.

458 THE WILD MAMMALS OF WISCONSIN Saltatorial — Adapted to jumping. Teat — The protruberance, i.e., the mamma Scansorial — Adapted to climbing. with nipple or at least with a nipple Scat — Fecal dropping. sucked during nursing. Scrotum — Sack of skin containing the testes Temporal — The squamosal and several ad- of mammals after birth. jacent minor bones comprise the tem- Seasonality — Temporal variation in climate poral complex; the temporalis muscles and availability of energy foods (or an help close the jaws having origin on the essential resource). squamosals and parietals. The so-called Sebaceous gland — A mammalian epidermal temporal bone is the squamosal. oil gland. Territoriality — The behavior of defending Sectorial — Adapted for shear or cutting. something, usually an area. Territory — Selenodont — With four crescentic cusps — An area defended. in Artiodactyla molars. Testis — The male sex organ that matures, Sign (sign also plural) — Scats, tracks, or in- emits sperm and produces testosterone. dications of a mammal. Testosterone — The hormone that influenc- Species — A population of mammals capa- es secondary sex characters, such as ble of freely interbreeding but reproduc- beard in man. tively isolated from other species. A tax- Tine — A pointed branch from a beam of an onomic category lower than the genus, antler. known by a binominal name. Torpor — A deep diurnal sleep characterized Speciation — Genetic and character differ- by only a slight drop in body tempera- entiation of two populations that might ture. Some workers use torpor and dor- or has resulted in evolution of new spe- mancy interchangeably. cies, which is usually encouraged by geo- Tragus — The slender, erect, spike-like pro- graphic isolation. cess from the lower part of the ear pin- Sperm storage — Delayed fertilization seen na in bats. in numerous bats and some marsupials. Transmigration of ova — The movement of a Spine — A stiff, sharp hair, a defensive fertilized egg from one horn of the uter- structure. us to the other, recognized by the cor- Succession — The sequence of change in spe- pus luteum being in the opposite ovary cies composition in a community as it of the cornu. passes through seral stages to a final sta- Tribosphenic — Trituberculate tooth, where ble climax stage. the three tubercles are so diminutive that Supraorbital ridge — A cranial prominence the tooth has little shear. In primitive over each orbit. mammals, the lower tricuspid tooth has Suture — An immovable joint extending lin- a talonid shelf bearing three small cusps. early (and vertically) between two bones. Tympanic bulla — See auditory bulla and Bulla. Sympatry — Where geographic ranges of Tympanic ring — Primitive, crescentic ecto- closely related mammals overlap. tympanic partially protects the middle Synonymy — A list of valid and invalid name ear. See endotympanic. combinations, with appropriate reference Type — Holotype — A specimen selected to included for each. represent a named kind of animal or plant. Sweat gland — A tubular gland of many mam- Underfur- The dense short fur that underlies mals that secretes a salty fluid on the skin the coarse guard hairs. to cool the body by evaporation. Ungulate — A hoofed animal. A general name Tarsal — Used as a noun, a bone of the for all hoofed mammals, which once had ankle. taxonomic use.

GLOSSARY OF MAMMALOGICAL TERMS 459 Unicuspidate (= unicuspate) tooth — A tooth Widths — Cranial, widest breadth of brain- with a single cusp, especially in the up- case. Zygomatic, widest breadth across per jaw of insectivores. Unicusp is more skull from one zygoma to the other. In- appropriate; unicuspid refers to lower terorbital, dorsal constriction across skull teeth, due to suffix -id. between orbits. Mastoid width, greatest Uropatagium — See interfemoral membrane. breadth across posterior part of skull, Velvet — The short fur on a developing antler. across mastoids. Venter — The lower or belly side. Wing-loading — Area of wings divided into Vestigial — An evolved small and variable the body weight. structure with impaired function. X chromosome — In XX combination causes Vibrissa — A whisker, an elongate special- female characters to develop. Y chromo- ized tactile hair. some — The XY pattern causes male Viviparous — Adapted to give birth to young, characters to develop. The Y is smaller, ova lacking shells. and has few genes. Volant — Although the word means aerial Zygomatic arch — An arch partially defend- the term is used for gliding forms. See ing the orbit comprised of an arm of the Patagium. squamosal, the jugal, the lacrimal, and Wisconsinan — Last major glacial advance, often the maxillary. Also called zygoma important in Wisconsin. (plural zygomata).

460 THE WILD MAMMALS OF WISCONSIN LITERATURE CITED AND Addison, E.M., M. J. Pybus, and J. J. Rietveld. BIBLIOGRAPHY 1978. Helminth and arthropod parasites of black bear, Ursus americanus, in central J. M., J. W. Mgt., WiWS, and WiAcadSciA&L = Ontario. Canadian J. Zool., 56: 2122-2126. J. Mammalogy, J. Wildlife Management, Advisory Committee of Immunization Practices. Wisconsin Wildlife Surveys, and Trans. 1991. Rabies prevention — United States. Wisconsin Acad. Sci., Arts and Letters Morbidity and Mortality Weekly Rpt., U.S. Dept. Health and Human Services, 40:1-19. Abbott H. C. and T. F. Quink. 1970. Ecology of Agassiz, G. R. 1913. Letters and recollections eastern white pine seed caches made by small of Alexander Agassiz... Houghton Mifflin forest mammals. Ecology, 51 :271-278. Co., Boston, N.Y. Ables, E. D. 1965. An exceptional fox move- Ainslie, W. B. 1983. Status, habitat preferenc- ment. J. M., 46: 102. es, and management of southwest Wiscon- Ables, E. D. 1969a. Activity studies of red fox- sin bats. M.S.Thesis, Univ. Wisconsin— es in southern Wisconsin. J. W. Mgt., 33: Stevens Point, 38 pp. 145-153. Aldous, S. E. 1941. Food habits of chipmunks. Ables, E. D. 1969b. Home-range studies of red J. M., 22: 18-24. foxes (Vulpes vulpes). J. M., 50: 108-120. Aldous, S. E. and J. Manweiler. 1942. The win- Ables, E. D. 1975. Ecology of the red fox in ter food habits of the short-tailed weasel in North America. Pp. 216-236, in M. Fox, northern Minnesota. J. M., 23: 250-255. Ed., The Wild Canids.... Van Nostrad Re- Aleksiuk, M. 1968. Scent-mound communica- inhold Co., New York. tion, terriroriality, and population regula- Adams, C. E. 1976. Measurements and charac- tion in beaver (Castor canadensis Kuhl). teristics of the fox squirrel, Sciurus niger J. M., 49: 759-762. rufiventer, home ranges. Amer. Midl. Nat., Aleksiuk, M. 1970. The function of the tail as a 95: 211-215. fat storage depot in the beaver (Castor ca- Adams, L. 1959. An analysis of a population of nadensis). J. M., 51: 145-148. snowshoe hares in northwestern Montana. Alexander, G.R. 1977. Food of vertebrate pred- Ecol. Monogr., 10:141-170. ators on trout waters in north central Lower Adams, R. A. 1992. Stages of development and Michigan. Michigan Acad. Sci, 10: 181-195. sequence of bone formation in the little Allen, D. L. 1938. Ecological studies on the ver- brown bat, Myotis lucifugus. J. M., 73: tebrate fauna of a 500-acre farm in Kalam- 160-167. azoo County, Michigan. Ecol. Monogr., 8: Adamsic, R. S. and L. B. Keith. 1978. Regional 348-436. movementsand mortality of great-horned Allen, D. L. 1939a. Michigan cottontails in win- owls in relation to snowshoe hare fluctua- ter. J. W. Mgt., 3: 307-322. tions. Canadian Field-Nat., 92: 228-234. Allen, D. L. 1939b. Winter habits of Michigan Adamsic, R. S., A. Wood, and L. B. Keith. skunks. J. W. Mgt., 3: 212-228. 1979.Demographic and dietary responses Allen, D. L. 1943. Michigan fox squirrel man- of red-tailed hawks during a snowshoe hare agement. Michigan Dept. Conservation, fluctuation. Canadian Field-Nat., 93:16-27. Game Div., 100:1-404. Addison, E. M., I. K. Barker, and D. B. Hunter. Allen, D. H. 1979. The wolves of Minong... 1987. Diseases and parasites of furbearers. Houghton Mifflin Co., Boston, 449 pp. In Wild furbearing mammals and conserva- Allen, D. L. and W. W. Shapton. 1942. An eco- tion in North America. by M. Novack, J. logical study of winter dens, with special Barker, M. Obbard and B. Malloch. eds. Min- reference to the eastern skunk. Ecology, istry of Nat. Resources, Ontario, Canada. 23: 59-68.

LITERATURE CITED AND BIBLIOGRAPHY 461 Allen, E. G. 1938. The habits and life history Anderson, R.M. 1946. Catalogue of Canadian of the eastern chipmunk, Tamias striatus Recent Mammals. Bull. Nat. Mus. Canada lysteri. New York State Mus. Bull. No. 102: v + 238 p. January 1947. 314, 22 pp. Anderson, S. H. 1982. Effects of the 1976 Seney Allen, G. M. 1933. Least weasel a circumboreal National Wildlife Refuge Wildfire on Wild- species. J. M., 14: 316-319. life and Wildlife Habitat. U. S. Dept. Interi- Alverson, W. S., D. M. Waller, and S. L. Sol- or, Fish and Wildlife Service, 146: 1-28. heim. 1988. Forests to deer: Edge effects Anderson, T. J. 1977. Population biology of the in northern Wisconsin. Conservation Bull., masked shrew, Sorex cinereus, in hard- 2: 348-358. wood forest areas of the McCormick Ex- Ambrose, H.W., III. 1973. An experimental perimental Forest, Marquette County, Mich- study of some factors affecting the spatial igan. M.S. Thesis, Northern Michigan and temporal activity of Microtus pennsyl- Univ., viii+75 pp. vanicus. J. M., 54: 79-110. Anon. 1995. Wisconsin’s biodiversity as a manage- Amin, O. M. 1974. Distribution and ecologi- ment issue. Dept. Nat. Resources. May 240 p. cal observations of wild mammals in Anthony, E. L .P., and T. H. Kunz. 1977. Feed- southeastern Wisconsin. WiAcadSciA&L, ing strategies of the little brown bat, Myo- 42: 311-326. tis lucifugus, in southern New Hampshire. Amin, O. M., and W. H. Thompson. 1974. Ar- Ecology, 58:775-786. boviral antibody survey of wild mammals in Applegate, V.C., and Predmore, H.E., Jr. 1947. southeastern Wisconsin. WiAcadSciA&L, Age classes and patterns of primeness in a 42:87-94. fall collection of muskrat pelts. J. W. Mgt. Amundson, T. and S. Marcquenski. 1985. Bay- 11: 324-330. lisascaris procyonis in Wisconsin raccoons. Archer. J., S J. Taft, and R. P. Thiel. 1986. Par- Offset report, Department of Natural Re- asites of wolves, Canis lupus, in Wiscon- sources, distributed but not published, Pp. sin, as determined from fecal examinations. 1-3 [4-7 unpaged, including 3 maps]. [Ex- Proc. Helminthol. Soc., 53: 290-291. cellent study of the prevalence of this par- Arlton, A.V. 1936. An ecological study of the asite in Wisconsin.] mole. J. M., 17:349-371. Andelt, W.F. and P.S. Gipson. 1979. Home Armitage, K. B. and K. S. Harris. 1982. Spatial range, activity, and daily movement of coy- patterning in sympatric populations of fox otes. J. W. Mgt., 43: 944-951. and gray squirrels. Amer. Midl. Nat., 108: Anderson, E. A. and A. Woolf. 1987. River ot- 389-397. ter habitat use in northwestern Illinois. Arnold, D.A. 1955. Status of Michigan timber Trans. Illinois Acad. Sci., 80: 107-114. wolves, 1955. Michigan Dept. Conserv., E. M. Anderson. 1987. A critical review and Game Div. Rept. No. 2062 (mimeo.), 2 pp. annotated bibliography of literature on the Arnold, D.A. 1956. Red foxes of Michigan. bobcat. Colorado Div. Wildife... Spec. Re- Michigan Dept. Conserv., 48 pp. port No. 62, 61 pp. Arnold, D.A. and R.D. Schofield. 1956. Home E. M.Anderson and M. J. Lovallo. 2004. Bob- range and dispersal of Michigan red foxes. caty and Lyx. Pp. 758-786, in Wild mam- Michigan Acad. Sci., 41:91-97. mals of North America. J. Chapman and Arthur, S. M., W. B. Krohn, and J. R. Gilbert. G. A. Feldhamer eds. Johns Hopkins Press. 1989a. Habitat use and diet of fishers. J. Anderson, R. M. 1942. Six additions to the list of W. Mgt., 53: 680-688. Quebec mammals with descriptions of four Arthur, S. M., W. B. Krohn, and J. R. Gilbert. new forms. Annual Report for 1941, Pro- 1989b. Home range characteristics of adult vancher Soc. Nat. Hist. of Canada, p. 31-43. fishers. J. W. Mgt., 53: 674-679.

462 THE WILD MAMMALS OF WISCONSIN Ashbrook, F.G. 1937. The breeding of fur ani- Ball, M. R. and J. Avise. 1992. Mitochondrial mals. Pp. 1379-1395 In Yearbook of Ag- DNA... The evolutionary significance of ric. U.S. Dept. Agric., Washington, D. C. subspecies. Auk., 109: 626-636. Atkeson, T. D. and R. L. Marchinton. 1982. Balliett, A. and S. J. Taft. 1976. Prevalence of Forehead glands in white-tailed deer. J. M., Dracunculus (Nematoda: Dracunculoidea) 63: 613-617. and Paragonimus kellicotti (Trematoda: Atkeson, T. D., R. L. Marchinton, and K. V. Troglotremaidae) is some Wisconsin mam- Miller. 1988. Vocalizations of white-tailed mals. Reports Fauna Flora Wisconsin, 14: 7. deer. Amer. Midl. Nat., 120: 194-200. Banfield, A.W.F. 1974. The mammals of Canada. Audubon, J. and J. Bachman. 1854. The vivip- Univ. Toronto Press, Toronto. xxiv+438 pp. arous quadrupeds of North America. 3 Banks, N. 1910. Ectoparasites of the rat. Pp. vols., New York. 67-85 In The rat and its relation to the Aumann, G.D. 1965. Microtine abundance and public health. U.S. Treas. Dept., Publ. Hlth. soil sodium levels. J. M., 46: 594-604. and Marine-Hosp. Serv., 254 pp. Avenoso, A.C. 1968. Selection and processing Barash, D. P. 1974. Mother-infant relations in of nuts by the flying squirrel, Glaucomys captive woodchucks (Marmota monax). volans. Ph.D Thesis, Univ. Florida, 126 pp. Animal Behavior, 22: 446-448. Bailey, B. 1893. The prairie ground squirrels or Barber, W. E. 1919. A beaver dam of huge pro- spermophiles of the Mississippi Valley. U.S. portions. Sci. American, 122(7):167. Dept. Agric., Div. Ornith. & Mamm., Bull. Barbour, R.W. and W.H. Davis. 1969. Bats of Amer- 4:1-69. ica. Univ. Kentucky Press, Lexington, 286 pp. Bailey, B. 1923. Meat-eating propensities of Barclay, R. M. R., P. A. Faure and D. R. Farr. some rodents of Minnesota. J. M., 4: 129. 1988. Roosting behavior and roost selection Bailey, B. 1929. Mammals of Sherburne Coun- by migrating silver-haired bats (Lasionycter- ty, Minnesota. J. M., 10: 153-164. is noctivagans). J. M., 69: 821-825. Bailey, B. 1937. The Maryland muskrat marsh- Barger, N. R. 1933. Bonaparte’s weasel (Mus- es. J. M., 18: 350-354. tela erminea bangsi). Wisconsin Cons. Bull., Bailey, J. A. 1969. Quantity of soft pellets pro- Maddison, 15: 39-40. duced by caged cottontails. J. W. Mgt., Barger, N. R. 1951. Prairie spotted skunk (Spilogale 33: 421. interrupta). Wis. Conserv. Bull. 16(5): 28. Baird, D. D., R. M. Timm, and G. E. Nordquist. Barnes, A. M. 1982. Surveillance and control of 1983. Reproduction in the Arctic shrew, bubonic plague in the United States. Symposia Sorex arcticus. J. M., 64: 298-301. of the Zoological Soc. London, 50: 237-270. Baird, Spencer F. 1857. Reports of explorations Barr, T.C., Jr. and R.M. Norton. 1965. Preda- and surveys to ascertain the most practica- tion on cave bats by the pilot black snake. ble and economical route for a railroad from J. M., 46: 672. the Mississippi River to the Pacific Ocean, Barr, T. R. B. 1963. Infectious diseases in the Vol. 8, pt. 1, Mammals, 757 pp. and 60 opossum. J. W. Mgt., 27: 53-71. [Review, pl. Actual date of publication 1858. chiefly S. Amer.] Baker, R.H. 1971. Nutritional strategies of my- Barrat, J., J. Blancou, C. Demantke, and Y. omorph rodents in North American grass- Gerard. 1985. b hemolytic streptococcal lands. J. M., 52: 800-805. infection in red foxes (Vulpes vulpes L.) in Baker, R. H. 1983. Michigan mammals. Ann France: the natural disease and experimen- Arbor, Michigan State Univ. Press. tal studies. J. Wildlife Diseases, 21:141-143. Baker, R. J. and J. L. Patton. 1967. Karyotypes Barron, R. J. 1981. Second record of the east- and karyotypic variation of North Ameri- ern pipistrelle Pipistrellus subflavus, in can vespertilionid bats. J. M., 48: 270-286. Michigan. The Jack-pine Warbler, 59: 68.

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486 THE WILD MAMMALS OF WISCONSIN Hoppe, R. T. 1979. Population dynamics of the Hsu, T. C. and K. Benirschke. 1967. An atlas of Michigan bobcat (Lynx rufus) with refer- mammalian chromosomes, Rattus. Vol. 1. ence to age structure and reproduction. Hsu, T. C. and K. Benirschke. 1967. An atlas Proceedings of the Bobcat Research Con- of mammalian chromosomes, Odocoileus ference, National Wildlife Federation Sci- virginianus. Vol. 1. entific and Technical Series, 6:111-115. Hsu, T. C. and K. Benirschke. 1968. An atlas Hornaday, H. T. 1904. The American natural of mammalian chromosomes, Mustela vi- history. New York. son. Vol. 2 Horner, B.E. 1954. Arboreal adaptations of Hsu, T. C. and K. Benirschke. 1970a. An atlas Peromyscus with special reference to use of mammalian chromosomes, Cryptotis of the tail. Univ. Michigan, Lab. Vert. Biol., parva. 4: 155. Contr. No. 61, 84 pp. Hsu, T. C. and K. Benirschke. 1970b. An atlas Hough, J. L. 1958. Geology of the Great Lakes. of mammalian chromosomes, Felis rufus. Univ. Illinois Press, Urbana. 4: 224. Houtcooper, W.C. 1971. Rodent seed supply Hsu, T.C., and K. Benirschke. 1971a. An atlas and burrows of Peromyscus in cultivated of mammalian chromosomes, Lepus fields. Indiana Acad. Sci., 81:384-389. townsendii., vol. 5, folio 211. Howard, W. E. 1949a. Dispersal, amount of Hsu, T.C. and K. Benirschke. 1971b. An atlas inbreeding, and longevity in a local popu- of mammalian chromosones, Microtus lation of prairie deer mice on the George ochrogaster. vol. 6, folio 272. Reserve, southern Michigan. niv. Michigan, Hsu, T. C. and K. Benirschke. 1971c. An atlas Lab. Ver. Biol. Contr., No. 43, 52 pp. of mammalian chromosomes, Mustela fre- Howard, W. E. 1949b. A means to distinguish nata. 5: 231. skulls of coyotes and domestic dogs. J. M., Hsu, T. C., and K. Berirschke. 1974. An atlas 30: 169-171. of mammalian chromosomes, Felis lynx. Howell, A. H. 1901. Revision of the skunks of the 8. [2 n = 38]. genus Chincha. N. Amer. Fauna, 20: 1-62. Huegel, C. N. 1979. Winter ecology of coyotes Howell, A.H. 1918. Revision of the American flying in northern Wisconsin. Univ. Wisconsin— squirrels. N. American Fauna No. 44, 64 pp. Madison, MS Thesis, 32. Howell, A. H. 1936. Revision of the North Huenecke, H.S., A.B. Erickson and W.H. Mar- American ground squirrels. N. American shall. 1958. Marsh gasses in muskrat hous- Fauna No. 56, 256 pp. es. J. W. Mgt., 22: 240-245. Hoy, P. R. 1853. The striped gopher, or prairie Hunsaker, D. 1977. Ecology of New World ground squirrel, of Wisconsin. Agric. Rept., marsupials. P. 95-156, in D. Hunsaker, ed., 1852. U.S.Patent Office, 2: 68-70. The biology of marsupials. Acad. Press, Hoy, P. R. 1875. On hibernation as exhibited New York. in the striped gopher. Proc. Amer. Assoc. Huff, J. N., and E.O. Price. 1968. Vocalizations Adv. Science, 24:148-150. of the least weasel, Mustela nivalis. J. M., Hoy, P. R. 1882. The larger wild mammals that 49: 548-550. have become extinct in Wisconsin. Humphrey, S.R., A.R. Richter and J. B. Cope. WiAcadSciA&L 5: 255-257. 1977. Summer habitat and ecology of the Hoyt, S. Y., and Hoyt, S.F. 1950. Gestation endangered Indiana bat, Myotis sodalis. J. period of the woodchuck, Marmota M., 58: 334-346. monax. J. M., 31: 454. Ilgin, N. A. 1941. Wolf-dog genetics. J. Genet- Hsu, T. C. and K. Benirschke. 1967. An atlas ics, 42: 359-414. of mammalian chromosomes, Mus muscu- Ingles, L. G. 1939. Observations on a nest of lus. Vol. 1. the long-tailed weasel. J. M., 20: 253-254.

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496 THE WILD MAMMALS OF WISCONSIN Long, C. A. 1975c. Rare and endangered mam- chickadee and white-breasted nuthatch. mals of Wisconsin. P. 237-245, In Cen- Wilson Bull., 94: 216-218. tennial Year Biol. Symp., Univ. Wisconsin- Long, C. A. 1982b. Phenotypic variation of size River Falls, 245 pp. and shape and the role of correlation in Long, C. A. 1975d. The distributional status and population morphology. Zhurnal Obschiei importance of bats of Wisconsin. P. 175- Biologii, 43: 360-365. 206, In Centennial Year Biol. Symp., Univ. Long, C. A 1985. Intricate sutures as fractal Wisconsin-River Falls, 245 pp. curves. J. Morphology, 185:285-295. Long, C. A. 1976a. Microtus ochrogaster mi- Long, C. A. 1986. Pleistocene caribou in central nor in Wisconsin. Univ. Wisconsin, Mus. Wisconsin. WisAcadSciA&L, 78: 12-13. Nat. Hist. Rpt., 11:1. Long, C. A. 1987a. A new subspecies of south- Long, C. A. 1976b. Notes on reproduction in ern bog lemming from two Lake Michigan pigmy shrews and observed ratios of mam- isles (Mammalia: Rodentia). Mammalia mae to body weights. Univ. Wisconsin, Mus. 51:324-326. Nat. Hist. Rpt., 11:5-6. Long, C. A. 1987b. Badger Lore. Wisconsin. Long, C. A. 1976c. Convergence and taxonomic Nat. Resources Mag. May/June, p. 29-30. relationships of some mustelids, especially Long, C. A. 1987c. Review of The Natural Histo- the badgers. Abstracts. Fifty-sixth Annual ry of Badgers, by E. Neal. Facts on File Publs., Meeting, Amer. Soc. Mammalogists, 24 New York & Oxford. J. W. Mgt., 51: 687. June Lubbock, Texas, p. 19. Long, C. A. 1988. Beaver. The World Book Long, C. A. 1977a. The occurrence, status and Encyclopedia, 2: 192-195. Based on prior importance of bats in Wisconsin with a key account. to the species. WisAcadSciA&L, 64:62-82. Long, C. A. 1988. Hare. The World Book Encyl- Long, C. A. 1977b. Evolution of mammalian clopedia, 9: 64. Snowshoe Hare, p. 17: 542. cheek pouches and a possible discontinu- Long, C. A. 1988. Muskrat. The World Book ous origin of a higher taxon (Geomyoidea). Encyclopedia, 13: 968. Amer. Midl. Nat. 110:1093-1097. Long, C. A. 1988. Woodchuck. The World Book Long, C. A. 1978a. An epiphysis on the scapu- Encyclopedia, 21: 394. la of the muskrat. Univ. Wisconsin, Mus. Long, C. A. 1990. Voles and bog lemmings of Nat. Hist. Rpt., 14:10-11. Wisconsin. WisAcadSciA&L, 78:87-110. Long, C. A. 1978b. Hartley Jackson and John Long, C. A. 1990. Weasel. The World Book Muir. Passenger Pigeon, p. 390. Enclyclo. Long, C. A. 1978c. Mammals of the islands of Long, C. A. 1990. Marten. The World Book Green Bay, Lake Michigan. Jack-Pine War- Encyclo. bler, 56::59-82. Long, C. A. 1992. Status and economic impor- Long, C. A. 1978d. A history of Recent bad- tance of the North American badger, gers of the world, with remarks on taxo- Taxidea taxus (Schreber). Newsl. and J. nomic problems in Mydaus and Melogale. IUCN Mustelid, Viverrid & Procyonid Spec. Reports on Fauna and Flora of Wisconsin, Group. 7: 4-7. 14: 1-6. Long, C. A. 1993a. Bivocal distraction nest-site Long, C. A. 1981. Provisional classification and display in the Red Squirrel, Tamiasciurus evolution of the badgers. Pp. 55-85, in J. hudsonicus, with comments on outlier nest- Chapman and D. Pursley, Proc. First World- ing and nesting behavior. Canadian Field- wide Furbearers Conf, 3 vols, Inst. Fish and Nat., 107:104-106. Wildlife Agencies, Frostburg, Maryland. Long, C. A. 1993b. Biogeography of the reptiles Long, C. A. 1982a. Comparison of the nest- and amphibians of the Lake Michigan Isles. site distraction displays of black-capped Bull. Chicago Herp. Soc., 28: 214-218.

LITERATURE CITED AND BIBLIOGRAPHY 497 Long, C. A. 1994. Fractal geometry in mam- dence in pigmentation patterns in mammals: All or none. Abstracts, Annual Meet- mals. J. M., 76: 937-939. ing, Amer. Soc. Mammalogists, Washing- Long, C. A., C. Becker, and A. Geier. 1976. ton, D. C., June. New records of rare shrews from the Lake Long, C. A. 1995a. Stone marten (Martes foi- Michigan Drainage Basin. Univ. Wisconsin, na) in southeast Wisconsin, U.S.A. News- Mus. Nat. Hist. Rpt., 11:2. letter and J. IUCN Mustelid, Viverrid & Long, C. A. and J. Captain. 1974. Investiga- Procyonid Species Group, 13: 14. tions on the sciurid manus. Part I. Some Long, C. A. 1995b. Wisconsin mammals: Some new taxonomic characters and their impor- tentative and taxonomic tales. Abstracts 75th tance in the classification of squirrels. Zeit. Annual Meeting Amer. Soc. Mammalogists, f. Saugetierkunde, 39:98-103. June 20-245 Burlington, Vt., p. 127. Long, C. A. and J. Captain. 1977. Investiga- Long, C. A. 1996. Ecological replacement of tions on the sciurid manus. Part II. Analysis the deer mouse, Peromyscus maniculatus, of functional complexes by morphological by the white-footed mouse, P. leucopus, integration and by coefficients of belong- in the Great Lakes Region. Canadian Field- ing. Zeit. f. Saugetierkunde, 42:214-221. Nat., 110: 271-277. Long, C. A. and F.A. Copes. 1968. Note on Long, C. A. 1999a. Sorex hoyi. Pp. 25-27, in the rate of dispersion of the opossum in D. Wilson and S. Ruff, eds. The Smithso- Wisconsin. Amer. Midl. Nat., 80: 283-284. nian Book of North American Mammals, Long, C. A. and T. Frank. 1968. Morphomet- Washington D. C. ric variation and function in the baculum, Long, C. A. 1999b. Taxidea taxus. Pp. 177- with comments on correlation of parts. J. 179, In D. Wilson and S. Ruff, eds. The M., 49: 32-43. Smithsonian Book of North American Long, C. A. and A. Hogan. 1988. Two inde- Mammals, Washington D.C. pendent loci for albinism in raccoons, Pro- Long, C. A. 2000. Book Review. Vaughan, T. cyon lotor. J. Heredity, 79:387-388. A., J. M. Ryan, and N. J. Czaplewski. Mam- Long, C. A. and T. Howard. 1976. Intraspecif- mals. 4th ed. Saunders College Publ. Co., ic overt fighting in the wild mink. Univ. Fort Worth, Texas. 565 pp. illus. J. M., Wisconsin, Mus. Nat. Hist. Rpts. 11:4-5. 81(3): 916-920. Long, C. A. and C. J. Jones. 1966. Variation Long, C. A. 2001. Extinction, eradication, and and frequency of occurrence of the bacu- reintroduction of some mammals in the lum in a population of Mexican free-tailed Wisconsin region, USA. 81 st Annual Meet- bats. Southwestern Nat., 11:290-295. ing American Mammalogists, June Missou- Long, C. A. and P. Kamensky. 1967. Osteo- la, Montana. Abstracts, No. 315; Interna- metric variation and function of the high- tional Theriological Congress, August, Sun speed wing of the free-tailed bat. Amer. City, SouthAfrica, Abstracts, No. 214. Midl. Nat. 77:452-461. Long, C. A. 2004. A principle of prolation in Long, C. A. and W. C. Kerfoot. 1963. Mam- biology. WSEAS Trans. Biol. and Biomed- malian remains from owl-pellets in eastern icine, 3(1): 311-315.[weasels] Wyoming. J. M.,44: 129-131. Long, C. A. 2006. Functional and physical extr- Long, C. A. and C.A. Killingley. 1983. The bad- tapolation relative to evolution of the mam- gers of the world. Charles C. Thomas Publ., malian erythrocyte. WSEAS Trans.Biol. and Springfield, Illinois, 404 pp. Biomedicine. 3(3): 195-203. Long, C. Alan. and C. A. Long. 1976. Some Long, C. A. and Joelle Affeldt Gehring. 1995. amphibians and reptiles collected on islands Valais-goat color pattern in a masked shrew in Green Bay, Lake Michigan. Jack-Pine (Sorex cinereus), and lack of size-depen- Warbler, 54: 54-58.

498 THE WILD MAMMALS OF WISCONSIN Long, C.A. and C.F. Long. 1965a. Dental abnor- Long, C.A., G. Zhang, T. George, and C.F. malities in North American badgers, genus Long. 1999 Bats. 80th Annual Meeting Taxidea. Kansas Acad. Sci., 68: 145-155. American Mammalogists, Durham, New Long, C.A. and C.F. Long. 1965b. Experienc- Hampshire, Abstracts, No. 246. es with adopted woodchucks. Defenders of Long, C.A., G. Zhang, T. George, and C.F. Wildlife News, 40: 59-61. Long. 1999. Physical theory, origin of Long, C.A. and C.F. Long. 1992. Some effects flight, and a synthesis proposed for birds. of land use on avian diversity in a Wiscon- J. Theoretical Biol., 224: 9-26. sin’s oak-pine savanna and riparian forest. Long, C. Alan. and C.A. Long. 1976. Some Passenger Pigeon, 54: 125-136. amphibians and reptiles collected on islands Long, C.A., C.F. Long, J. Knops, and D.H. in Green Bay, Lake Michigan. Jack-Pine Matulionis. 1965. Reproduction in the dick- Warbler, 54: 54-58. cissel. Wilson Bull., 77: 251-256. Long, C.F. 1971. Common grackles prey on Long, C.A. and J.E. Long. 1986. Mass drown- big brown bat. Wilson Bull., 83: 196. ing of gray squirrels of Washington Island, Longley, W.H. 1962. Movements of red fox . J. Lake Michigan. The Jack-pine Warbler, 64: M., 43: 107. 21-22. Longley, W.H. 1963a. Minnesota gray and fox Long, C.A. and J.E. Long. 1988. Southern bog squirrels. Amer. Midl. Nat., 69: 82-98. lemming, Synaptomys cooperi, new to is- Longley, W.H. and J.B. Moyle. 1963. The lands in Lake Michigan. Canadian Field- beaver in Minnesota. Minnesota Dept. Nat., 102: 64-65. Cons., Game and Fish Tech. Bull., No. Long, C.A. and J.E. Long. 1992. Fractal dimen- 6, 87 pp. sions and wave forms. Acta Anat. 12 p. Lopez-Rivera, N. In Prep. Population indices of Long, C.A. and J.E. Long. 1993. Discriminant small mammals in central Wisconsin. Per- analysis of geographic variation in long- sonal correspondence. tailed deer mice from northern Wisconsin Lord, R.D., Jr. 1958. The importance of juve- and Upper Michigan. WisAcadSciA&L, 81: nile breeding to the annual cottontail crop. 107-121. Trans. Twenty-third N. Amer. Wildl. Conf., Long, C.A. and R.G. Severson. 1969. Geo- p. 269-276. graphical variation in the big brown bat in Lord, R.D. 1959. The lens as an indicator of the north-central United States. J. M., age in cottontail rabbits. J. W. Mgt. 28: 50:621-624. 358-360. Long, C.A. and L.R. Shirek. 1970. Variation and Lord, R.D. 1960. Litter size and latitude in north correlation in the genital bones of ranch mink. American mammals. Amr. Midl. Nat. 64: Zeit. f. Saugethierkunde, 35: 252-255. 488-499. Long, C.A. and D. Smart. 1976. Variation and Lord, R.D., Jr. 1961. The lens as an indicator correlation of the os cordis and heart in of age in the gray fox. J. M., 42: 109-111. Odocoileus virginianus (Artiodactyla). Tex- Lotz, J.M. and W.F. Font. 1983. Review of as J. Sci., 27 155-161. the Lecithodendriidae (Trematoda) from Long, C.A. and C.J. Yahnke. 2007. Post-gla- Eptesicus fuscus in Wisconsin and Min- cial caribou and elk-moose in Late Pleis- nesota. Proc. Helmintholog. Soc. Wash- tocene-Holocene sediments in central Wis- ington, 50: 83-102. consin. Quart. Res., submitted. Lotze, J.-H. 1985. The raccoon (Procyon lo- Long, C.A., G. Zhang, and T. George. 2002. tor) on St. Catherines Island, Georgia. IV. Physical and evolutionary problems in take- Comparisons of home ranges determined off runs of bipedal winged vertebrates. Ar- by livetrapping and radiotracking. Amer. chaeopteryx. 20: 63-71. Mus. Novit., No. 2664, 25 pp.

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526 THE WILD MAMMALS OF WISCONSIN Zajac, A.M. and J.F. Williams. 1980. Infection Zimmerman, E.G. 1965. A comparison of hab- with Schistosomatium douthitii (Fam. itat and food of two species of Microtus. Schistosomatidae) in the meadow vole (Mi- J. M., 46: 605-612. crotus pennsylvanicus) in Michigan. J. Zimmerman, E.G. 1972. Growth and age de- Parasitol., 66: 366-367. termination in the thirteen-lined ground Zakrzewski, R.J. 1985. The fossil record. In R.H. squirrel Spermophilus tridecemlineatus. Tamarin, ed., Biology of the New World Amer. Midl. Nat., 87: 314-325. Microtus, Amer. Soc. Mammalogists, Spec. Zimmerman, G.D. 1974. Cooperative nursing Publ. No. 8. behavior observed in Spermophilus tride- Zeckmeister, M.T. and N.F. Payne. 1998. Effects cemlineatus (Mitchill). J. M., 55: 680-681. of trapping on colony density, structure, and Zimmerman, J.H. 1991. The landscape and the reproduction of a beaver population unex- birds. Pp. 35-90, in S. D. Robbins, Jr. Wis- ploited for 19 years. WiAcadSciA&L, 86: consin Birdlife... Univ. Wisconsin Press, 281-291. Madison, Wisconsin. Zegers, D.A. and J.F. Merritt. 1988. Adapta- Zimmerman, W.J. 1970. The epizoology of tri- tions of Peromyscus for winter survival in chianiasis [trichinosis] in wildlife. J. Wildlife an Appalachian montane forest. J. M., 69: Diseases 6: 329-334. 516-523. Zimmerman, W.J. 1971. Trichinosis in wildlife. Pp. Zielinski, W.J. 1978. A supernumerary and 127-139, in Parasitic diseases of wild mam- anomalous tooth in Peromyscus leuco- mals. Iowa State Univ. Press, Ames, Iowa. pus. Univ. Wisconsin Mus. Nat. Hist. Zimmerman, W.J. 1977. Trichinosis in bears in Rpt., 14: 8-9. western and north-central United States. Zielinski, W.J. 1988. The influence of daily vari- Amer J. Epidemiol., 106: 167-171. ation in foraging cost on the activity of small Zimny, M.L. 1965. Thirteen-lined ground squir- carnivores. Animal Behaviour, 36: 239-249 rels born in captivity. J. M., 46: 521-522. Ziman, E. 1976. On the regulation of pack size Zinsser, H. 1935. Rats, lice and history. Little in wolves. Zeit. f. tierpsychol. 40: 300-341. Brown and Co., Boston, Mass. 228 pp.

LITERATURE CITED AND BIBLIOGRAPHY 527 APPENDIX A Longevities of Some Mammals That Occur in Wisconsin (After Crandall, 1964)

Species Years Didelphis virginiana 3-4 Blarina brevicauda 2 Lepus americanus 3-8+ Sciurus carolinensis 9-15 Sciurus niger 9 Tamiasciurus hudsonicus 5-9 Marmota monax 7-9 Spermophilus franklinii 7 S. tridecemlineatus 8 Tamias striatus 3-7 Glaucomys volans 7 Castor canadensis 13 Urocyon cinereoargenteus 10 Vulpes vulpes 10-12 Canis latrans 12-15 Canis lupus 12-16 Ursus americanus 19-25 Procyon lotor 15 Martes americana 13-18 Neovison vison 5-10+ Taxidea taxus 13-2 Mephitis mephitis 6-7 Lontra canadensis 13-19 Lynx canadensis 11 Lynx rufus 15 Alces alces 8 (M); 9 (F)

528 THE WILD MAMMALS OF WISCONSIN APPENDIX B English and Metric Measurements

Inches (in) Millimeters (mm) 1 25.4 2 50.8 12 305 36 914.5 Feet Millimeters 1 305 3 914.5 Yards 1 914.5 1 + 3 3/8 in 1000 (= 1 m) 100 (= 300 ft) 91.45 meters Miles (mi) Kilometers 1 1.61

Ounces (oz) Grams (g) 129 16 (1 lb) 453.6 0.04 1 Pounds (lbs) Kilograms (Kg) 5 2.27 25 11.3 100 45.4 2.2 1 Fahrenheit (F) Celsius (C) 32o (freezing point) 0o 100o 37o 212o (boiling point) 100o Acres Hectares (ha) 1 (= 43,560 ft2 0.4 =4,047 m2) 2.46 1 Quarts Liters (l) 1 0.946 1.057 1 (=1,000 cc)

APPENDIX B 529 APPENDIX C scales are available. The sign for female or Preparation of the Mammal Study male sex is written in the label’s upper left Skin with Label and Skull corner. This label will be tied eventually to the specimen’s right hind foot, and care must be taken that it is not tied to some other spec- LIST OF SUPPLIES imen. On the backside of the label the species binomen may be written, but it is not es- z Skin labels and skull labels sential. A skull label is also prepared for the z Permanent black ink and permanent skull, and on it should be written the collec- ink pen tor’s name (not initials) and his field number. z Millimeter rule, preferably a flexible The sex is also useful. plastic rule z Sewing needles and stout thread 2. Skinning the mammal z Scalpel or single-edge razor blade After measuring the standard measurements, z Forceps with slender tines the body is removed inside out from its skin z Scissors with sharp points through a small abdominal incision, and the z Long-fibered cotton, white (not sur- appendages are freed from the skin in this gical cotton) order: hind leg, hind leg, tail, foreleg, fore- z Pan or pie-tin leg, and skull. The hind legs are cut free at z White cornmeal or fine hardwood the ankles, or metarsals in furbearers, the fore- sawdust legs at the wrists, or metacarpals in furbear- z Borax to be powdered lightly on the ers. The tail (vertebrae) are pulled out with internal skin of the specimen (never care so that the tail skin is NOT turned wrong on the skull or skeleton, which is side out, even if one must hold the skin tight- cleaned by dermestids, that would ly at its base as the tail vertebrae are with- suffer from the borax) drawn. With a razor or scalpel cut free the z Field Scales, weighing in grams. ear pinnae, eyes, and nose cartilage, careful- z Notebook of good quality for per- ly separating the skin. Use a sharp blade to manent record cut the cartilage from the nasal bones, and z Non-rusting wire pulled out straight do not cut them. The anterior corner of each and cut with sharp ends eye must be cut free with great care or the z Good wire cutters to cut the wire eyelids will be torn open enormously. Now lengths sharp the carcass is in one hand and the skin is in z A tackle box or container for these the other. Clean blood and fat away from the supplies to carry in the field. skin’s inside surface. Roll it up to keep it from drying or place it in a plastic bag.

STAGES OF PREPARATION 3. Skull or skeleton If preparing only the skull without the post- 1. Measuring and label preparation cranial skeleton attached to it, cut the neck, The permanent label written with permanent through the cervical vertebrae. With forceps black ink has on it the locality of capture, the thrust the thread of the skull label through date, the collector’s name (in full) and his cat- the jaws and out the mouth and tie the thread alog number (from his field catalog and jour- so that the skull is labeled with name and field nal), and the standard measurements. These number of the collector. The skull should be are total, tail, hind foot, and ear lengths, sup- dried somewhat, not hard, and it is ready for plemented with the body weight in grams if cleaning in the dermestid colony of the mu-

530 THE WILD MAMMALS OF WISCONSIN seum. By removing the viscera and most of out to the tail tip. Leave bare extensions of the flesh from the postcranial carcass, the all four wires alongside the cotton body to entire skeleton attached to the skull may be support the feet and tail. Then insert sharp cleaned in the dermestid colony. This skele- wires into the forelegs (along the cotton body, ton is, of course, minus the foot bones which inside the skin) piercing them into the feet. are still in the skin. Before the skeleton dries These should also have extensions of wire left somewhat, roll it up like a ball, and loop thread alongside the cotton body for support. Bits of around it to keep all the bones together. Also, inserted (with forceps) cotton may advanta- the baculum found in males, should be left in gaeously puff out all four limbs. Pull incision the prepuce dangling on the skin, divorced shut, wrapping skin around body and wire carefully from the penis as the tail is removed extensions. Sew up the incision with a few from the skin. The baculum is important for stitches. Slap or clap the stuffed specimen into identification and should always be preserved. a normal position with feet and tail straight back, the forefeet tucked under the chin, and 4. Providing a new body the hind feet bracing the tail from either side. A new body is made up of loose-fibered cot- The skin label was tied on the right hind foot. ton to resemble the carcass, by making a single U-shaped roll into a cylinder. Pinch the 5. Pinning and drying anterior end and thrust it through the abdom- The study skin is pinned on a drying board, inal incision within the skin, which of course even a cardboard box, so that the forefeet has fur outward, and push or pull the pinched are under the chin, the eyes pinched shut, cotton end with forceps into the head as far forward as possible (to the mouth and nose). Tear off extra cotton at the other end so that one can insert that end into the rump area. If one has not made the incision great, it is possible to insert this cotton end into the posteri- or pocket of skin near the tail. Wires are never put in before the cotton body, insert the sharp end of a straight wire into each hind leg piercing the foot. Insert two wires into the empty tail skin to puff it out full,, one wire

App-3-2App-3-2. Measuring the tail from the pelvis to its fleshy App-3-1. Skin label and skull label. The stain was tip (not including the hairs). Cutting free the left hind foot. App-3-1. Skin label and skull label. blood. Cornmeal should blot up blood. Remove all muscles. Sewing up the incision. After Hall.

APPENDIX C 531 the ears straightened up, the mouth closed, Safeguard the specimen from diverse and the hind feet and tail projected straight dangers (pets, people, elements, insects) un- behind. The body should be full. In a few days til a curator can catalog it into a museum and the skin will dry, provided one properly re- afford perpetual care. Ten water shrews in moved flesh and fat from the skin. Most pro- the U.S. National Museum were prepared fessionals use ten pins per specimen (4 feet, over 100 years ago by President Theodore 4 per tail (2 and 2), 2 straightening the hind Roosevelt, and they are in perfect condition legs). Usually the skull is pinned alongside each with skins, skulls, and data written on the la- specimen to dry. bels. Specimens are forever.

532 THE WILD MAMMALS OF WISCONSIN APPENDIX D Dental Records*

Didelphis virginiana. Lepus townsendii.

Lepus americanus. Juvenile Lepus americanus.

Sylvilagus floridanus. Juvenile Sylvilagus floridanus.

* Photos by S. Sepsenwol and C. Long. Nikon Coolpix 4500 digital camera. The coin is 10 mm in diameter.

APPENDIX D 533 Marmota monax. Spermophilus tridecemlineatus.

Spermophilus franklinii. Eutamias minimus.

Tamias striatus. Tamiasciurus hudsonicus.

534 THE WILD MAMMALS OF WISCONSIN Sciurus carolinensis. Sciurus niger.

Glaucomys sabrinus. Glaucomys volans.

Castor canadensis. Geomys bursarius.

APPENDIX D 535 Microtus pennsylvanicus. Ondatra zibethicus.

Erythizon dorsatum. Canis latrans.

Vulpes vulpes. Urocyon cinereoargenteus.

536 THE WILD MAMMALS OF WISCONSIN Ursus americanus. Procyon loter.

Mustela nivalis. Neovison vison.

Mustela erminea. Mustela frenata.

APPENDIX D 537 Mephitis mephitis. Lontra canadensis.

Taxidea taxus. Lynx rufus.

Odocoileus virginianus. Lynx canadensis .

538 THE WILD MAMMALS OF WISCONSIN INDEX TO SCIENTIFIC AND Canis latrans (See coyote) 24, 40, 50, 52, 65, VERNACULAR NAMES OF WILD 323-324, 326, 426, 528, 536 Spec. 3683. MAMMALS OF WISCONSIN Canis lupus (See timber wolf = gray wolf) 19, 40, 50, 65, 323-324, 330-333, 337, 528. Alces alces (See American moose) 51, 60, 65, Casteroides ohioensis 56-57. 439, 450, 452, 528. Castor canadensis (See North American beaver) AMERICAN MINK (See Neovison vison) 6, 12, 19, 22, 40, 49-50, 57, 64, 155, 220-222, 14, 31, 36, 46, 49, 65, 80, 96, 124, 172, 226, 228, 528, 535 Spec. 7152. 174, 191, 226, 228, 248, 278, 289, 296- Cervalces scotti 55, 58. 298, 308, 314, 342, 364, 366-370, 372, Cervus elaphus (See North American elk) 12, 356-376, 379, 381, 392. 14, 22, 24, 50, 60, 65, 439. AMERICAN OR NORTH AMERICAN MOOSE Cervus Nippon Sika deer 61-62, 439. (See Alces alces), 6, 31, 35-36, 53, 55, CINEREOUS SHREW (see Sorex cinereus) 64, 86. 58, 60, 65, 67, 335, 431, 438-440, 446, Clethrionomys gapperi (See red-backed vole) 37, 450-452, color plates. 51, 56-57, 64, 261-264, 373. AMERICAN OR PINE MARTEN (See Martes Condylura cristata (See Star-nosed mole) 38, americana) 43, 49, 59, 65, 367, 369-370, 40, 50, 52, 64, 76, 78-79, 277, 373. . 372, 375-379, 399. COYOTE (See Canis latrans) 6, 14, 28, 49, 65, AMERICAN PORCUPINE (See Erethizon dor- 92, 148, 154, 184, 101, 260, 289, 321- satum) 6, 14, 26, 42, 50, 53, 65-66, 156, 325, 327-331, 340, 342-343, 346, 364, 315-320, 326-327, 335, 369, 374-375, 391, 426, 446. 378, color plates. Cryptotis parva (See least shrew) 18-19, 31, 38, ARCTIC SHREW (See Sorex arcticus) 53, 64, 40, 48-50, 56, 59, 64, 75, 85, 101-102, 86, 89-90, 92. 106-108, 342. BADGER See Taxidea taxus. Dama dama Fallow deer 62, 439. BEAVER See Castor canadensis. 457. Dicrostonyx torquatus 56. BIG BROWN BAT (See Eptesicus fuscus) 64, Didelphis virginiana (See opossum) 48-49, 51, 113, 118, 131-132, 135, 141. 56, 64, 67, 69, 528, 533 Spec. 6734. Bison bison 15, 48, 60, 65, 336, 412, 438- EASTERN CHIPMUNK (See Tamias striatus) 439. 38, 49, 64, 157, 176, 179-180, 183-184, Bison latrifrons 56-57. 198, 372, color plates. Bison occidentalis 55-57. EASTERN COTTONTAIL (See Sylvilagus flo- BLACK BEAR (See Ursus americanus), 6, 44, ridanus) 14, 38, 44, 50, 64, 73, 144, 147, 49, 65, 226, 318, 321, 351-352, 354-357, 150-151, 153-155, 342, 349, 363, 372, 359, 446, color plates. 397, 437. Blarina brevicauda (see Blarina and short-tailed EASTERN MOLE (See Scalopus aquaticus) 31, shrew) 36, 40-43, 49-50, 52, 64, 75, 77, 76, 81-82, 84, color plates. 85, 89, 96-97, 101-105, 235, 254, 264, EASTERN SPOTTED SKUNK (See Spilogale 266, 282, 284, 342, 372-373, 396, 426. putorius) 53, 65, 367, 403, 410-413. BOBCAT (See Lynx rufus) 12, 14, 36, 49, 53, Elephas primigenius 55, 57-58. 65, 74, 88, 104, 254, 184, 191, 199, 213, Eptesicus fuscus (See big brown bat) 64, 110, 224, 226, 265, 289, 297, 315, 322, 364, 113, 118, 130-131. 402, 408, 417, 427-430, 432-438, 443, Erethizon dorsatum (See American porcupine) color plates. 50, 54, 65, 316-317, 373, 536 Spec. 6737. CANADA LYNX (See Lynx canadensis) 29, 31, ERMINE (See Mustela erminea) 38, 53, 65, 368, 34-36, 49, 50, 53, 65, 191, 226, 373, 383, 387-391, 395, 537 Spec. 5731, co- 402, 408, 427-435. lor plates.

INDEX 539 Eutamias minimus (See least chipmunk) 24, 40, Lasiurus borealis (See red bat) 49-50, 64, 110, 45, 51, 64, 157, 174-180, 534 Spec. 2895. 112-113, 132, 135-137, 139, 141-142, EVENING BAT (See Nycticeius humeralis) 64- Lasiurus cinereus (See hoary bat) 64, 111-112, 65, 112, 130-131. 138-142. Felis canadensis See Lynx canadensis. LEAST CHIPMUNK (See Eutamias minimus) Felis catus 14, 16, 61. 45, 53, 64, 157, 175-177, 179, 186, co- Felis concolor 12, 19, 24, 50, 59, 65. lor plates. Felis rufus See Lynx rufus. LEAST SHREW (See Cryptotis parva) 31, 53, FISHER (See Martes pennanti) 10, 19, 44, 49, 59, 64-65, 85, 106, 108, color plates. 59, 65, 161, 169. LEAST WEASEL (See Mustela nivalis) 14, 53, 65, FOREST DEER MOUSE (see Peromyscus ma- 278, 321, 368, 382-383, 385-387, 395-396. niculatus) 18, 52-53, 155, 240, 250, 252. Lepus americanus (See snowshoe hare) 38, 40, FOX SQUIRREL (See Sciurus niger) 53, 64, 43, 51-52, 57, 64, 144, 146-150, 154, 157, 166, 193-194, 203, 205-209, 363, 372, 396, 431, 528, 533 Spec. 1813. color plates. Lepus townsendii (See white-tailed jack rabbit) 48, FRANKLIN’S GROUND SQUIRREL (See Spermo- 50, 53, 59, 64, 144-147, 533 Spec. 1021. philus franklinii) 53, 64-65, 157, 170-174. LITTLE BROWN BAT (See Myotis lucifugus) Geomys bursarius (see plains pocket gopher) 19, 64, 113, 117-118, 120, 141. 24, 27, 50, 56, 64, 156, 229-231, 233, LONG-TAILED WEASEL (See Mustela frenata) 235, 424, 535 Spec. 700. 28, 65, 92, 191, 368, 383, 388, 392-393, GEORGIAN PIPISTRELLE (See Perimyotis sub- 395-398. flavus) 64, 112, 127-129, 140, color plates. Lontra canadensis (See river otter) 19, 40, 50, Glaucomys sabrinus (See Northern flying squir- 65, 367, 414, 416, 528, 538 Spec. 8421. rel) 37, 40, 45, 51-52, 54, 64, 157, 209- Lutra Canadensis (See Lontra canadensis. 219, 373, 431, 535 Spec. 4927. Lynx canadensis (See Canada lynx) 40, 51, 65, Glaucomys volans (see Southern flying squirrel) 150, 428, 430-431, 433, 528, 538 Spec. 36, 43, 45, 48, 50, 56, 59, 654, 157, 210- 7868. 211, 213-217-219, 528, 535 Spec. 1457. Lynx lynx See Lynx canadensis. GRAY FOX (See Urocyon cinereoargenteus) 6, 13- Lynx rufus (See bobcat) 19, 50, 52, 65, 433- 14, 65, 278, 321-322, 339-342, 344, 420. 434, 436, 438, 528, 538 Spec. 1882. GRAY SQUIRREL (see Sciurus carolinensis) 52, Mammut americanum 55, 57-58. 64, 157-158,171, 187, 193-194, 196- Marmota monax (See woodchuck) 40, 43, 45, 200, 202-203, 208. 50, 64, 155, 157-160, 162, 164, 426, GRAY WOLF (See also timber wolf and Canis 528, 534 Spec. 7144. lupus) 213, 289, 330-331, 333, 408. Martes americana (See American marten) 51, GROUNDHOG 64, 157-159. See woodchuck 59, 65, 367, 369-370, 372, 375-377, 528. and Marmota monax Martes foina (See stone marten) 12, 22, 65, 367, Gulo gulo 12, 22, 51, 57, 59, 65, 368. 378. HOARY BAT (See Lasiurus cinereus) 64, 112, Martes pennanti (See fisher) 44, 51, 59, 65, 150, 138, 140-141. 318, 322, 367-378, 391, 399, 402, 408. HOUSE MOUSE (see Mus musculus) 12, 59, MASKED SHREW (see cinereous shrew and 61, 155-156, 235, 242, 298, 301-303. Sorex cinereus) 44, 49, 51, 64, 86-89, 97, INDIANA MYOTIS (See Myotis sodalis) 41, 64- 100, 372, 431, color plates. 65, 72, 113, 121-122, 124. MEADOW JUMPING MOUSE (See Zapus hud- KEEN’S MYOTIS (See Myotis keenii) 64, 114-115. sonius) 65, 305-306, 308, 311, 372. Lasionycteris noctivagans (See silver-haired bat) MEADOW VOLE (see Microtus pennyslvanicus) 110 64, 112-113, 124-127, 141-142. 42-43, 49, 64, 92, 104, 261, 274, 280-285.

540 THE WILD MAMMALS OF WISCONSIN Mephitis mephitis (See striped skunk) 19, 50- NORTH AMERICAN BEAVER (See Castor ca- 51, 57, 65, 110, 367, 403-406, 409, 412, nadensis) 6, 13-14, 17, 26, 41, 49, 51-52, 528, 538 Spec. 292. 64, 66, 94-95, 155-156, 185, 219-228, Microsorex hoyi (see Sorex hoyi and Sorex 291-292, 316, 322, 335, 342, 346, 356, (Microsore) hoyi and pygmy shrew) 22, 24, 372, 417, 419, 431, 437, color plates. 27, 51, 64, 85-86, 92, 96-97, 100-101. NORTH AMERICAN ELK or wapiti (See Cervus Microtus ochrogaster See Pitymys ochrogaster elaphus)12, 24, 49, 55, 57-58, 60-61, Microtus pennyslvanicus (See meadow vole) 14, 318, 439-440, 447, 450-451. 40, 64, 104, 150, 155, 261, 266, 268- NORTHERN FLYING SQUIRREL (See Glauco- 269, 271-272, 274, 278, 280-283, 285- mys sabrinus) 31, 45, 53, 64, 157, 210- 287, 327, 536 Spec. 3830. 211, 214. Microtus pinetorum See Pitymys pinetorum NORWAY RAT (see Rattus norvegicus) 12, 59, Microtus xanthognathus 56, 62. 63, 65, 155-156, 184, 235, 298-300, 303. MINK See American mink. Nycticeius humeralis (See evening bat) 18, 22, MOOSE See American moose. 40, 49-50, 64, 110, 112-113, 115, 130- Mus musculus (See house mouse) 16, 19, 59, 131, 142. 61, 65, 155, 235, 237, 242-243, 257, Odocoileus virginianus (See white-tailed deer) 298, 301-303. 24, 50, 65, 295, 373, 439-441, 538 Spec. MUSKRAT (See Ondatra zibethicus) 10, 14, 36, 3682. 38, 41-42, 49, 65, 73, 155-156, 228, 261, Ondatra zibethicus (See muskrat) 40, 49-50, 65, 291-298, 322, 335, 346, 363, 372, 399, 418. 150, 292-294, 373, 536 Spec. 3077. Mustela erminea (See ermine) 38, 40, 51, 65, OPOSSUM (See Didelphis virginiana) 11, 13- 368, 381-382, 385, 387-390, 392, 537 14, 31, 43, 45-46, 51, 53, 61, 64, 66-75, Spec. 5731. 134, 136, 154, 278, 315, 322, 342, 363, Mustela frenata (See long-tailed weasel) 40, 50, 437, 458, color plates. 65, 368, 382, 384-385, 388, 392-394, OTTER (See Lontra canadensis). 396, 537 Spec. 1017. Perimyotis subflavus (See Georgian pipistrelle) Mustela nivalis (See least weasel) 40, 51, 65, 49-50, 57, 64, 112-113, 127-130, 142. 321, 368, 381-386, 388, 537 Spec. 8043. Peromyscus leucopus (See white-footed mouse) Mustela vison See Neovison vison. 19, 38, 40-41, 43-45, 48, 50-52, 56, 64, Myotis keenii (See Keen’s Myotis) 40, 50, 52, 104, 235, 240-254, 256- 260, 327, 392. 64, 110, 113-117, 141. Peromyscus maniculatus (see Forest deer mouse Myotis lucifugus (See little brown bat) 37, 40, and Prairie deer mouse) 12, 22, 36, 43, 50, 64, 110-111, 113-115, 117, 119, 121- 45, 50, 52, 57, 64, 155, 236-237, 241- 124, 142. 242, 246, 250-253, 255, 259. Myotis sodalis (See Indiana bat or Indiana Myotis) Peromyscus maniculatus bairdii (see Prairie deer 22, 49, 64, 113, 115, 117, 121-124, 142. mouse) 12, 27, 36-38, 40, 43, 45, 50, 64, Napaeozapus insignis (See woodland jumping 236-237, 241-242, 244, 249, 252-253, mouse) 40, 50, 65, 305, 311-312. 255, 257-259. Neovison vison (See American mink) 40, 50, Peromyscus maniculatus gracilis 48, 251-252 65, 150, 297, 368, 392, 398-404, 403, (see Peromyscus maniculatus maniculatus). 408, 418, 528, 537 Spec. 3636. Phenacomys intermedius 56-57. NORTH AMERICAN BADGER (See Taxidea Phenacomys ungava 57. taxus) 6, 13-14, 26, 41, 49, 65, 67, 168, PINE VOLE (see woodland vole and Pitymys pi- 174, 190-191, 220, 232-233, 325, 335, netorum) 53, 64, 104, 264, 267, 274-275, 342, 360, 366-367, 391, 404, 420-421, 277-279, 284. 423-427, color plates. Pipistrellus sp. (See Perimyotis) 127, 130.

INDEX 541 Pitymys ochrogaster (= Microtus ochrogaster) RIVER OTTER (See Lontra canadensis) 6, 40, (See prairie vole and also P. o. minor) 22, 49, 53, 65, 154, 227-228, 366-367, 403, 38, 40, 43-44, 48-49, 50, 53, 57, 64, 414-421, color plates. 150, 235, 261, 266-269, 273-275, 278, Scalopus aquaticus (See prairie or eastern mole) 281, 287. 38, 50, 56, 64, 76-78, 81, 83-84, 396. Pitymys ochrogaster minor (see Pitymys ochro- Sciurus carolinensis (see gray squirrel) 19, 22, gaster and prairie vole) 12, 38, 53, 132, 36, 43, 50, 64, 150, 157, 193-195, 203- 268-269, 271-274, 315. 204, 208, 528, 535 Spec. 5664. Pitymys pinetorum (see Woodland vole and Pine Sciurus niger (See fox squirrel)43, 48, 50, 59, vole) 22, 24, 40, 48-51, 56, 64, 266-268, 64. 157-158, 166, 196, 203-205, 208- 274-276, 279, 286. 209, 528, 535 Spec. 4932. PLAINS POCKET GOPHER (see Geomys bur- SHORT-TAILED SHREW (See Blarina brevicau- sarius) 6, 27, 29, 31, 38, 41, 53, 64, 66, da) 85, 101-102, 104, 372 . 156, 229-230, 232-234, 396. SILVER-HAIRED BAT (See Lasioncyteris noc- Platygonus compressus 56-57. tivagans) 110, 64, 112, 124-125. PORCUPINE See American porcupine. SNOWSHOE HARE (See Lepus americanus) PRAIRIE DEER MOUSE (see Peromyscus ma- 38, 53, 64, 144, 147-148, 150, 195 , 326- niculatus bairdii) 12, 18, 36, 38, 53, 241, 327, 335, 375, 418, 427, 432. 257, 259. Sorex arcticus (See Arctic shrew) 40, 51, 56- PRAIRIE DOG 61. 57, 64, 75, 85-86, 89-92, 97. PRAIRIE MOLE (See (See Scalopus aquaticus) Sorex cinereus (See masked shrew and cinere- 31, 38, 53, 64, 76-77, 81-82. ous shrew) 36, 40, 42, 44, 51, 64, 84, 85- PRAIRIE VOLE (see Pitymys ochrogaster) 12, 87, 90, 92-93, 97-98, 104-105. 38, 43-44, 53, 64, 262, 268-269, 271-274. Sorex fumeus 62, 85. Procyon lotor (See raccoon) 19, 50, 57, 65, Sorex hoyi and Sorex (Microsorex) hoyi (also see 340, 359-362, 373, 528, 537 Spec. 2186. Microsorex hoyi and Pygmy shrew) 12, 22, PYGMY SHREW (See Sorex hoyi) 49, 53, 64, 24, 57, 64, 85-86, 92, 96-99, 100-101. 85, 97, 98, 100. Sorex palustris (See water shrew) 24, 40, 43, RACCOON (See Procyon lotor) 6, 13-15, 42, 49, 48, 51, 56-57, 64, 85, 93-95, 97, 372. 65, 67, 154, 184, 208, 260, 278, 322, 335, SOUTHERN BOG LEMMING (See Synaptomys 340, 346-347, 359-366, 378, 405, 420. cooperi) 36, 38, 52, 64, 156, 261, 264, Rangifer tarandus 12, 22, 51, 56-57,65. 268, 275, 285-287, 289-291, 335. Rattus norvegicus (see Norway rat) 12, 16, 59, SOUTHERN FLYING SQUIRREL (See Glauco- 63, 65, 155, 235, 299-301, 390. mys volans) 38, 41, 64, 157, 211, 214- RED BAT (See Lasiurus borealis) 49, 53, 64, 215, 217-218, color plates. 112-113, 135-136, 138, 141. Spermophilus franklinii (See Franklin’s ground RED FOX (See Vulpes vulpes) 6, 13, 26, 14, squirrel) 50, 64, 157, 165, 170-173, 528, 49, 65, 108, 184, 187, 191, 200, 260, 534 Spec. 318. 278, 321-322, 339-349, 364, 373, 402. Spermophilus tridecemlineatus (See 13-lined RED SQUIRREL (See Tamiasciurus hudsonicus) ground squirrel or thirteen-lined ground 53, 64, 154, 157-158, 176, 184, 186-190, squirrel) 26-27, 37, 40, 45, 48-50, 57, 64, 192, 194, 198, 200, 335, 426. 164-165, 169-171, 244, 528, 534 Spec. RED-BACKED VOLE (see Clethrionomys gap- 4392. peri) 38, 53, 64, 261-262, 264-266, 275. Spilogale putorius (See Eastern spotted skunk) Reithrodontomys megalotis (see Western har- 22, 48, 50, 65, 367, 410-413. vest mouse) 24, 38, 43, 50-51, 56, 64, 235- STAR-NOSED MOLE (See Condylura cristata) 236, 238, 243. 38, 53, 64, 67, 76-77, 80, color plates.

542 THE WILD MAMMALS OF WISCONSIN STONE MARTEN (See Martes foina) 12, 367, Urocyon cinereoargenteus (See gray fox) 19, 378-381, color plates. 22, 24, 50-52, 65, 321, 339-340, 345, STRIPED SKUNK (See Mephitis mephitis) 13- 347, 528, 536 Spec. 1048. 14, 49, 65, 72, 260, 315, 322, 361, 365, Ursus americanus (See black bear) 40, 50, 57, 367, 372, 404-405, 407-409, color plates. 65, 321, 351-353, 528, 537 Spec. 2770. Sylvilagus floridanus (See Eastern cottontail) 38, Vulpes vulpes (See red fox) 40, 50, 57, 65, 150, 40, 43-45, 48, 50, 59, 64, 143-144, 147, 321, 339-340, 343, 345, 350, 528, 536 150-152, 373, 390, 396, 401, 426, 533 Spec. 1815. Spec. 4200. WATER SHREW (See Sorex palustris) 49, 53, Symbos and Ovibos 57. 64, 85, 90, 93-94, 372, 532. Synaptomys borealis 56. WESTERN HARVEST MOUSE (see Reithrodon- Synaptomys cooperi (See Southern bog lem- tomys megalotis) 24, 49, 51, 53, 64, 156, ming) 19, 24, 36, 40, 50-52, 57, 64, 261, 236-237, 239-240, 243, 302-303. 279, 284, 286, 288-291. WHITE-FOOTED MOUSE (see Peromyscus leu- 13-LINED GROUND SQUIRREL or Thirteen- copus) 36, 38, 41, 52, 64, 104, 242, 244- lined Ground Squirrel (See Spermophilus 246, 249-250, 257, 283, 372, 544. tridecemlineatus) 26-27, 45, 53, 64, 157- WHITE-TAILED DEER or whitetail (See Odocoi- 158, 164-169, 171-172, 229-230, color leus virginianus) 6, 13, 16, 41-42, 45, 49, plates. 62, 65, 224, 295, 327, 332, 335, 431, 437- Tamias striatus (See Eastern chipmunk) 13, 22, 444, 447, 449, 451-452, color plates. 24, 36, 43, 50, 56-57, 64, 157, 175-176, WHITE-TAILED JACK RABBIT (See Lepus 179-182, 185-186, 249, 528, 534 Spec. townsendii) 49, 53, 59, 64-65, 144-146. 1626. WOODCHUCK (See Marmota monax) 6, 44, Tamiasciurus hudsonicus (See red squirrel) 19, 64, 155, 157-159, 161-163, 325, 335, 36, 40, 43, 51, 57, 64, 157, 186-188, 342, 372, 407, 418, color plates. 190, 192-193, 200, 231, 373, 396, 426, WOODLAND JUMPING MOUSE (See Napaeoza- 528, 534 Spec. 2607. pus insignis) 38, 63, 65, 305, 311-312, 315. Taxidea taxus 24, 38, 40, 50, 65, 367, 421- WOODLAND VOLE (see pine vole and Pitymys 422, 528, 538 Spec. 4031. pinetorum) 261, 268, 274-275, 279, co- Thomomys talpoides 56-57. lor plates. TIMBER WOLF (see also gray wolf and Canis Zapus hudsonius (See meadow jumping mouse) 36, lupus) 49, 65, 315, 323-324, 330-333, 44, 51, 56, 65, 303, 305, 307, 309-312. 337, 339, 357, color plates. Zapus princeps 56.

INDEX 543 The white-footed mouse. Drawn by Anna Stryke.

544 THE WILD MAMMALS OF WISCONSIN TEST YOUR KNOWLEDGE! Every species has a fascinating story for Wisconsin. 8 4 6 2 7 5 3 1 10

15 16 17 18 19

1. Opossum 19. Ermine 2. Star-nosed Mole 20. Striped Skunk 3. Eastern Mole 21. River Otter 4. Masked or Cinereus Shrew 22. American Badger 5. Least Shrew 23. Bobcat 6. Georgian Pipistrelle 24. White-tailed Deer 7. Woodchuck (= Groundhog) 25. American Moose 8. Least Chipmunk. By Tyler Long 9. Eastern Chipmunk 10. Thirteen-lined Ground Squirrel 11. Fox Squirrel 12. Southern Flying Squirrel 13. Beaver 14. Woodland or Pine Vole 15. Porcupine 16. Timber or Gray Wolf 17. Black Bear 18. Stone Marten 20

21 22

23 22

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