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2005

Prairie Dog Empire: A Saga of the Shortgrass

Paul A. Johnsgard University of Nebraska - Lincoln, [email protected]

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Johnsgard, Paul A., " Empire: A Saga of the Shortgrass Prairie" (2005). University of Nebraska Press -- Sample Books and Chapters. 6. https://digitalcommons.unl.edu/unpresssamples/6

This Article is brought to you for free and open access by the University of Nebraska Press at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in University of Nebraska Press -- Sample Books and Chapters by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. 1 2 3 4 5 6 7 8 9 10 Prairie Dog Empire 11 [First Page] 12 13 [-1], (1) 14 15 Lines: 0 to 22 16 17 ——— 18 * 388.16002pt PgVar ——— 19 Normal Page 20 * PgEnds: PageBreak 21 22 23 [-1], (1) 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

Kim—UofNPress/Pagei//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 Other Titles by Paul A. Johnsgard 2 Published by the University of Nebraska Press 3 4 Lewis and Clark on the : A Natural History 5 The Nature of Nebraska: Ecology and Biodiversity 6 This Fragile Land: A Natural History of the Nebraska Sandhills 7 Crane Music: A Natural History of American Cranes 8 Birds of the 9 Diving Birds of North America 10 Those of the Gray Wind: The Sandhill Cranes 11 The Platte: Channels in Time 12 The Grouse of the World 13 The Plovers, Sandpipers, and Snipes of the World [-2], (2) 14 A Guide to North American Waterfowl 15 Birds of the Great Plains: Breeding Species and Their Distribution Lines: 22 to 70 16 Song of the North Wind: A Story of the Snow Goose 17 The Bird Decoy: An American Art Form ——— * 18 Waterfowl of North America 243.822pt PgVar ——— 19 North American Game Birds of Upland and Shoreline Normal Page 20 Grouse and Quails of North America * PgEnds: PageBreak 21 Waterfowl: Their Biology and Natural History 22 23 [-2], (2) 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

Kim—UofNPress/Pageii//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 2 3 4 5 6 7 Prairie Dog Empire 8 9 10 A Saga of the 11 12 Shortgrass Prairie 13 [-3], (3) 14 15 Lines: 70 to 102 16 17 paul a. johnsgard ——— 18 * 62.822pt PgVar ——— 19 Normal Page 20 * PgEnds: PageBreak 21 22 23 [-3], (3) 24 25 26 27 28 29 30 31 32 33 34 35 University of Nebraska Press, Lincoln and London 36 37 38 39 40

Kim—UofNPress/Pageiii/ /Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 © 2005 by the Board of 2 Regents of the University of Nebraska 3 All rights reserved 4 Manufactured in the 5 United States of America ⅜ϱ 6 Library of Congress Cata- 7 loging-in-Publication 8 Data 9 Johnsgard, Paul A. Prairie dog empire : a saga 10 of the shortgrass prairie / 11 Paul A. Johnsgard. 12 p. cm. Includes bibliographical [-4], (4) 13 references and index. 14 isbn 0-8032-2604-7 15 (hardcover : alk. paper) Lines: 102 to 179 16 1. Prairie ecology – Great Plains. 2. Prairie animals – ——— 17 Great Plains. I. Title. 18 qh104.5.g73j64 2005 * 205.976pt PgVar ——— 19 578.74'4'0978 –dc22 2004024696 Normal Page 20 Set in Minion by Kim * PgEnds: PageBreak 21 Essman. 22 Designed by Dika Eckersley. 23 Printed by Thomson- [-4], (4) 24 Shore, Inc. 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

Kim—UofNPress/Pageiv//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 2 3 4 5 6 7 8 9 10 Dedicated to the people and groups who have long and valiantly worked to 11 preserve our native heritage 12 13 [-5], (5) 14 15 Lines: 179 to 192 16 17 ——— 18 * 374.58002pt PgVar ——— 19 Normal Page 20 * PgEnds: PageBreak 21 22 23 [-5], (5) 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

Kim—UofNPress/Pagev//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 2 3 4 5 6 7 8 9 10 11 12 13 [-6], (6) 14 15 Lines: 192 to 194 16 17 ——— 18 0.0pt PgVar ——— 19 Normal Page 20 PgEnds: T X 21 E 22 23 [-6], (6) 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

Kim—UofNPress/Pagevi//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 2 3 4 5 Contents 6 7 8 9 10 List of Maps viii 11 List of Illustrations viii 12 List of Tables ix 13 [-7], (7) Preface xi 14 15 1. The Western Shortgrass Prairie: A Brief History 1 Lines: 194 to 267 16 2. A Buffalo Nation: Its Death and Rebirth 12 17 3. Prairie Dogs and the American West: Little Towns on the Prairie 28 ——— 18 4. Ferrets, Badgers, Bobcats, and : 7.18979pt PgVar ——— 19 Coping with Dangerous Neighbors 47 Normal Page 20 5. Free-loaders and Hangers-on: * PgEnds: PageBreak 21 The Rewards and Dangers of City Life 61 22 23 6. Other High Plains Wildlife: Born on the American 79 [-7], (7) 24 7. The High Plains Raptors: Death Rides on Silent Wings 95 25 8. The Varmint and Predator Wars: 26 It’s Finally Almost Quiet on the Western Front 106 27 9. Taming the Great American Desert: 28 Hardscrabble Times at the Fringes of Nowhere 118 29 30 10.Theusfs, blm, and bia: How the West Was Lost 129 31 11. The Great Plains Grassland Ecosystem: Can It Be Saved? 146 32 appendix 1. 33 A Guide to National , Reservations, 34 and Nature Preserves on the High Plains 151 35 appendix 2. 36 Scientific Names of Animals in the Text 193 37 38 References 199 39 Index 237 40

Kim—UofNPress/Pagevii/ /Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 maps 2 1. Ecoregions of the Great Plains 9 3 2. Bird Conservation Regions of the High Plains 10 4 3. Biological regions of the Great Plains 11 5 4. Distribution of 16 6 5. Current distribution of the five prairie dog species 32 7 6. Distribution of 62 8 7. Distribution of and kit fox 69 9 8. Distribution of 81 10 9. Breeding distribution of 86 11 12 illustrations [-8], (8) 13 1. Bison, males head-to-head fighting 24 14 2. Bison, male bellowing 24 15 3. Bison, male following estrus female 26 Lines: 267 to 408 16 4. Bison, male tongue-extension display 26 ——— 17 5. Bison, female nursing calf 27 18 9.87637pt PgVar 6. Adult black-tailed prairie dog 30 ——— 19 7. Utah prairie dog, adult 34 Normal Page 20 8. Gunnison’s prairie dog, adult 35 PgEnds: TEX 21 9. White-tailed prairie dog, adults in upright postures 38 22 10. Black-tailed prairie dog, adult jump-yip display 43 23 11. Black-tailed prairie dog, adult and young “kissing” 44 [-8], (8) 24 12. Black-tailed prairie dog, female nursing 45 25 13. Black-tailed prairie dog, crouching and sunbathing 48 26 14. Black-tailed prairie dog, alarm calling 48 27 15. Adult black-footed ferret 50 28 16. Adult black-footed ferret 50 29 17. Adult badger 52 30 18. Adult bobcat 55 31 19. Adult 57 32 20. Adult burrowing owl 64 33 21. Adult swift fox 67 34 22. Hispid pocket mouse, Ord’s kangaroo rat, mouse, and 35 grasshopper mouse 73 36 23. McCown’s and chestnut-collared longspurs, lark bunting, 37 horned lark, and mountain plover 76 38 24. horned lizard, tiger salamander, and ornate box turtle 78 39 25. Adult male pronghorn 80 40

Kim—UofNPress/Pageviii/ /Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 26. Spotted, thirteen-lined, and Richardson’s ground squirrels 84 2 27. Long-billed curlew, marbled godwit, upland sandpiper, and 3 mountain plover 87 4 28. Horned lark, upland sandpiper, marbled godwit, and 5 mountain plover 88 6 29. McCown’s longspur, chestnut-collared longspur, and lark bunting 93 7 30. Adult golden eagle 96 8 31. Golden eagle and prairie dog 98 9 32. and prairie dog 99 10 33. Adult prairie falcon 102 11 34. Bison, adult male 126 12 35. Bison, adult female 126 13 [-9], (9) 14 tables 15 1. Estimated historic and recent U.S. black-tailed prairie dog populations 33 Lines: 408 to 414 16 2. Recent acreage and population estimates for three prairie dog species 36 ——— 17 3. Summer prairie dog and burrow densities 37 18 * 160.92148pt PgVar 4. Coyote foods based on stomach analyses 58 ——— 19 5. Relative food item occurrences in scats of swift foxes from five states 71 Normal Page 20 6. Vertebrate associates of black-tailed prairie dogs 74 * PgEnds: PageBreak 21 7. Coyotes reported killed by sport hunters and taken in usda’s 22 Wildlife Services Program during 2000–2001 in five High Plains states 108 23 8. Estimated black-tailed prairie dog acreages, 1870 and 1998 112 [-9], (9) 24 9. Recent prairie dog poisoning statistics for three Great Plains states 113 25 10. Some estimated annual kills of prairie dogs by recreational shooting 115 26 11. Mixed-grass and shortgrass prairie acreage estimates in the Great Plains 122 27 12. blm, national grassland, and homestead acreages in the Great Plains 132 28 29 30 31 32 33 34 35 36 37 38 39 40

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Kim—UofNPress/Pagex//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 2 3 4 5 Preface 6 7 8 9 10 In early July of 2002 I attended a hearing at the Lincoln headquarters of Ne- 11 braska’s Game and Parks Commission. The main agenda item at this hearing was 12 whether the State of Nebraska would begin to take steps toward conserving the 13 black-tailed prairie dog. As a result of a 1998 petition from the National Wildlife [-11], (11) 14 Federation, followed up later by one from the Biodiversity Legal Foundation, 15 the species had been legally proposed as a candidate for listing as nationally Lines: 416 to 437 16 threatened. Nebraska is one of eleven states within the species’ current range 17 and had been previously notified by the U.S. Fish and Wildlife Service that ——— 18 such a listing, with all its associated required conservation measures, could be 0.0pt PgVar ——— 19 averted only if these states individually and collectively began concerted efforts Normal Page 20 to begin preserving the species. PgEnds: T X 21 An hour or more of oral testimony by conservationists, biologists, and rep- E 22 resentatives of various environmental groups ensued, which uniformly favored 23 initiating a modest conservation program, including some supportive testi- [-11], (11) 24 mony from biologists of the Game and Parks Commission itself. Then a group 25 of ranchers who had been bused in from across the state took the floor, con- 26 demning the prairie dog with all the usual vituperation that has been associated 27 with rancher–prairie dog relationships of the past century or more. At the 28 end of the meeting the commissioners unanimously voted not to provide any 29 conservation efforts whatsoever toward prairie dogs and furthermore voted to 30 terminate all state-supported research on the species’ status and biology that 31 was then being undertaken or planned by the commission’s staff biologists. 32 The ban on future prairie dog studies was later rescinded, but no steps toward 33 producing a conservation management plan were taken. 34 As a result of that meeting, I decided that enough interest exists in prairie 35 dogs, including strongly held attitudes both pro and con, to warrant a book 36 centered on this controversial animal. Such a book would also describe and 37 document the semiarid grassland ecosystem of the western plains within which 38 the prairie dog evolved and in which it has historically played a pivotal, or 39 “keystone,” ecological role. In many ways, by the late 1900s the black-tailed 40 prairie dog was providing a reprise of the sad history of the North American

Kim—UofNPress/Pagexi//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 bison, whose genocidal destruction during the late 1800s spelled the end of the 2 American frontier and likewise brought an end of the entire culture that had 3 been represented by the bison-dependent Dakotas and other Native Americans 4 of the western plains. Now, roughly a century later, many ranchers who had 5 displaced the Native Americans were themselves being threatened, as a com- 6 bined result of prolonged , long-depressed markets, and ranges 7 so badly overgrazed and weed infested that most of the small operators could 8 not provide the numbers of livestock needed to make a living. 9 This book is a result of these experiences and related concerns. As an ornithol- 10 ogist I have become increasingly worried about the decline and often regional 11 disappearance of the burrowing owl, a commensal associate of the prairie dog. 12 I also fear for the ferruginous hawk, whose nest sites and winter distributions in 13 the Great Plains and American Southwest often center on prairie dog “towns,” [-12], (12) 14 and for the tiny swift fox, an agile and beautiful little carnivore that once also 15 was common but now is largely confined to areas of prairie dog towns. The swift Lines: 437 to 443 16 fox is presently so rare (as of 2004 it is a federal candidate species for a listing 17 as threatened) that I have yet to see one in the wild. By the time I began my ——— 18 writing, it was already perhaps too late to save the black-footed ferret, another 0.0pt PgVar ——— 19 prairie dog–dependent carnivore that is already federally listed as endangered Normal Page 20 and that is surviving only by the thinnest of life-support threads. The ferret, one PgEnds: T X 21 of America’s rarest mammals, was one of the first mammal species to be placed E 22 on the nationally endangered list. Nevertheless, the North was 23 also once reduced to perhaps no more than a hundred or so animals surviving [-12], (12) 24 in the wild. Yet the species has rebounded and now numbers several hundred 25 thousand animals, mostly in captive herds. 26 All of these symbols of the western plains have close ecological ties to prairie 27 dogs. If prairie dogs are not to be saved, we have little hope of preserving some 28 of the others either, to say nothing of affecting the dozens of additional animals 29 that are also part of and variously dependent on the prairie dog–bison–buffalo 30 grass ecosystem of the American plains. 31 I have relied on the help of many people in assembling the information 32 needed for this book. Among them are Craig Knowles of FaunaWest Wildlife 33 Consultants, who provided me with several valuable unpublished reports on 34 the status of three species of prairie dogs. Robert Luce, interstate coordina- 35 tor, Prairie Dog Conservation Team, also provided me with some important 36 references and an advance copy of the proceedings of a Conservation Team 37 workshop meeting held in late 2003. Tyler Sutton of the Great Plains Con- 38 servation Alliance sent me various unpublished reports on prairie dogs and 39 plans for establishing a high plains ecological preserve south of the Black Hills. 40 “Buffalo” Bruce MacIntosh and Gerald Jasmer provided me with valuable un-

xii preface

Kim—UofNPress/Pagexii/ /Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 published materials on the prairie dog and the swift fox. Al Steuter of the Nature 2 Conservancy’s Niobrara Valley Preserve loaned me materials on bison biology 3 and bison rearing. Joe Truett of the Turner Biodiversity Fund also provided me 4 with unpublished information on the foundation’s important conservation and 5 restoration programs, and Francis Moul gave me a manuscript copy of some of 6 his graduate research on the history of the national grasslands. Barbara Voeltz 7 of the Nebraska Game and Parks Commission helped me with Internet searches 8 of the extensive biological literature of the western plains grasslands. I also wish 9 to thank the three anonymous persons who reviewed the manuscript for the 10 University of Nebraska Press. 11 I decided that before I finished writing I needed to visit some of the remaining 12 natural grassland regions within the vast Great Plains range of the black-tailed 13 prairie dog, and especially the major National Grassland reserves that are at the [-13], (13) 14 heart of the species’ remaining range. For advice or help with fieldwork, provid- 15 ing reference materials, and other assistance, I am indebted to Doug Backlund, Lines: 443 to 447 16 Linda Brown, Jackie Canterbury, Bob Gress, Josef Kren, Bob Luce, Al Steuter, 17 Tom Shane, and Scott Wendt. All of these people helped me understand the ——— 18 often neglected and invariably sad histories of all of the historic residents of our * 49.58002pt PgVar ——— 19 western American , from prairie dogs, , and bison to our Native Normal Page 20 American kin, whose lives and fortunes were fatally and inexorably intertwined * PgEnds: PageBreak 21 with the prairie. 22 In a broader sense I owe a special debt of gratitude to the organizations that 23 are fighting to preserve the prairie dog and its associated prairie ecosystem as [-13], (13) 24 part of our American heritage and that have helped to document their status. 25 These groups include such diverse public interest organizations as Biodiversity 26 Associates, the Biodiversity Legal Foundation, the Center for Native Ecosystems, 27 the Conservation Alliance for the Great Plains, Forest Guardians, the Fund for 28 Animals, the Humane Society, the National Wildlife Federation, The Nature 29 Conservancy, the Prairie Dog Coalition, the Predator Conservation Alliance, 30 Rocky Mountain Animal Defense, and the Turner Biodiversity and Endangered 31 Species Funds. Without the documented information and related historic mate- 32 rials these groups have assembled, and without their legal maneuvering against 33 entrenched federal agencies and powerful opposing private interests, this book 34 might not have been written. Without their efforts, not enough of the prairie 35 dog ecosystem would be left to bother to preserve, and the story of the prairie 36 dog would have to be told only in the past tense. 37 38 39 40

preface xiii

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Kim—UofNPress/Pagexvi/ /Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 2 3 1 4 5 The Western Shortgrass Prairie 6 7 A Brief History 8 9 10 There was nothing but land. Not a country at all, but the materials out of which countries are made. – Willa Cather, My Antonia 11 [First Page] 12 13 Like the animals and plants that now live on it, the land composing the surface [1], (1) 14 and substrate of the North American Great Plains nearly all came from some- 15 place else. Some of the region’s parts were carried by westerly winds as volcanic Lines: 0 to 56 16 ash from mountains a thousand or more miles to the west, while others were 17 blown in as dust-sized particles from areas up to several hundred miles to the ——— 18 northwest. Most of what geographers call the Great Plains was carried in and 0.0pt PgVar ——— 19 deposited by and streams originating in the Rocky Mountains or was Normal Page 20 randomly strewn as glacial-carried materials from the north. Only in a few ar- PgEnds: T X 21 eas, such as the igneous and metamorphic Precambrian core of ’s E 22 Black Hills and the 300 million-year-old (Paleozoic) Wichita Mountains of 23 , are the remnants of the earth’s early violent history exposed in the [1], (1) 24 form of ancient and eroded mountains (Trimble 1990). 25 In its simplest form, the present-day Great Plains region may be thought of 26 as a poorly constructed tabletop that is slightly tilted downward toward the 27 east-southeast. Its western limits are formed by the towering Front Range of 28 the Rocky Mountains, cresting at the Continental Divide, and grading into 29 their outlying piedmonts, which end indecisively at about 5,000 to 6,000 feet 30 of elevation. From there the Great Plains take over, gradually losing altitude 31 eastwardly. At roughly 1,000 feet of elevation, the Great Plains merge invisibly 32 with the Central Lowlands, a region massively shaped by glaciers at the north- 33 eastern end and forming a shallow basin made up of the Missouri, Mississippi, 34 and Ohio valleys toward the southeast. These river valleys were cut and 35 repeatedly reformed by the immense meltwaters of several glaciations, carving 36 broad, fertile valleys as they slowly traveled to deposit their silty cargoes along 37 the way or dump them into the Gulf of Mexico. 38 The southern parts of the Great Plains become increasingly flat and arid, 39 including the treeless and so-called Staked Plains of the Texas Panhandle and 40 eastern , a land so flat and barren that early explorers reputedly

Kim—UofNPress/Page1//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 drove tall wooden stakes into it at intervals to keep from becoming lost. The 2 southern terminus of the Great Plains is decisively marked in New Mexico by the 3 Mescalero Escarpment and the associated Pecos Valley, and in Texas the Plains 4 are also abruptly terminated southwardly by the Edwards Plateau. The northern 5 end of the Great Plains disappears quietly somewhere near Great Slave Lake 6 in ’s vast transcontinental coniferous forest, an enormous area largely 7 scraped down by glacial action nearly to ancient bedrock, the Precambrian 8 Shield of the earth’s earliest crust. 9 Within the Great Plains region, the higher, more arid western portions that 10 now support only shortgrass and mixed-grass prairies dominated by perennial 11 herbs of low to middle stature can be geologically and biologically distinguished 12 as the High Plains. Geologically speaking,the northern edge of the High Plains is [2], (2) 13 the Pine Ridge Escarpment of northwestern Nebraska and southwestern South 14 Dakota. From there south, most of the High Plains consists of a nearly flat 15 surface, covered by sandy materials called the Ogallala formation, a layer up to Lines: 56 to 68 16 several hundred feet thick of water-transported materials that were deposited 17 ——— in Pliocene times. The upper part of this layer is commonly a thick zone of 18 13.0pt PgVar very hard carbonates, producing a “caprock” of rocklike material (caliche) that ——— 19 is almost impervious to water. This zone, not very evident in the northern Normal Page 20 part of western Nebraska’s western plains, becomes thicker southwardly and PgEnds: T X 21 E is up to 30 feet thick at the southern end of the western plains in Texas. The 22 conspicuous Caprock Escarpment, extending in a general north-northeast to 23 [2], (2) south-southwest direction from western Oklahoma through the Texas Panhan- 24 dle, marks the southeastern edge of the caprock zone and provides a convenient 25 26 southern bookend to the High Plains (Thornbury 1965). 27 North of the Pine Ridge Escarpment and the nearby Black Hills is the Mis- 28 souri Plateau of eastern , northeastern , and the western 29 Dakotas, and the Missouri River valley itself. Eastward and northward from the 30 Missouri River in the Dakotas is a fertile region of glacial drift and till deposited 31 by the last glaciation, forming a substrate for the biologically rich prairie 32 wetlands region of the northern plains. Although technically not a part of the 33 High Plains as they have been geologically defined, these portions of the west- 34 ern and central Dakotas, eastern Montana and Wyoming, and the southernmost 35 prairie-covered portions of and share most of the same 36 semiarid and grassland-adapted plant and animal life that is now found in 37 the more southern parts of the Great Plains, including a , the 38 black-tailed prairie dog. For this book’s purposes these areas are included in the 39 general region here informally recognized as the western plains, or “shortgrass 40 prairie ecosystem.”

2 the western shortgrass prairie

Kim—UofNPress/Page2//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 a geological time line 2 3 To understand the current overall patterns of plant and animal distributions in 4 the western plains, one must think backward a few million years. Until about 70 5 million years ago, all of what is now the Great Plains lay beneath a vast inland 6 sea. At its bottom were sediments up to 10,000 feet deep. Around 70 million 7 years ago the North American continent began a gradual uplift, progressively 8 exposing the sea bottom and producing a series of gentle basins and arches on 9 the rising landscape. However, more extensive uplifting occurred in the Black 10 Hills region, and the Rocky Mountains to the west began their much more 11 extensive expansions, largely through folding and faulting. As rapidly as the 12 western mountains rose, rivers formed to erode their surface layers, carrying 13 sediments away to the more easterly vegetated plains, where a variety of plant- [3], (3) 14 eating and carnivorous dinosaurs were dining on what would – by about 65 15 million years ago – became their final meals. The river sediments preserved the Lines: 68 to 78 16 fossil remains of such impressive but now defunct animals as Triceratops and 17 Tyrannosaurus. ——— 18 About 50 million years ago volcanic activities in mountains far to the west 0.0pt PgVar ——— 19 added additional wind-carried materials, even as abundant tropical vegetation Normal Page 20 still flourished along the edges of the retreating seas. In places like the northern PgEnds: T X 21 plains these lush forests of tropical evergreen vegetation were eventually buried E 22 to become the vast beds of lignite and coal that are being extracted today from 23 Wyoming, Montana, and the western Dakotas. [3], (3) 24 Between about 45 million and 35 million years ago a pause in mountain- 25 building occurred. Gradually drier and cooler climates and less-forestlike plant 26 communities began to develop, only to be interrupted by a renewal of volcanism 27 and mountain building near the end of this period. More river-related erosion 28 brought new layers of sediments to the Plains, including the White River sedi- 29 ments that compose much of South Dakota’s scenic badlands. By now the land 30 east of the Rocky Mountains was mostly covered by a vast grassland and many 31 large grazing mammals. Early horses and camels had displaced the dinosaurs 32 and other dominant reptiles of the late Mesozoic era. Fossil remnants of some 33 of these high-toothed grazing mammals were preserved in such places as Agate 34 Fossil Beds National Monument in western Nebraska, among sediments more 35 than 20 million years old. 36 Between 25 and 5 million years ago the Great Plains were affected by periodic 37 phases of mountain building and volcanism, each episode adding layers of new 38 sediments and windblown debris to the older materials progressively interred 39 below. The oldest deposits of sediments occurred in the northern parts of the 40 Great Plains, while more recent deposits of Oligocene, Miocene, and Pliocene

the western shortgrass prairie 3

Kim—UofNPress/Page3//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 vintages extended progressively farther east and south, eventually reaching the 2 southern edge of the present-day Great Plains. A rather late surge of volcanism 3 in the western mountains occurred about 9 million years ago. This activity 4 suffocated herds of rhinos and other large mammals under a cloud of volcanic 5 dust, providing the fossilized base for what is now known as Ashfall Fossil Beds 6 State Historical Park, in north-central Nebraska. By 5 million years ago the 7 entire High Plains region was becoming a vast, eastwardly sloping tableland 8 similar to its present shape, with numerous rivers – including the predecessors 9 of the Missouri, the Platte, the Cimarron, and the Pecos – all cutting downward 10 through the soft surface materials near the mountains and redepositing them 11 farther downstream. 12 About 5 million years ago a new period of uplift occurred in western North 13 America, causing new cycles of down-cutting and subsequent deposition, with [4], (4) 14 rivers such as the Missouri eroding down to depths only a few hundred feet 15 higher than their current elevations. Prior to the Pleistocene glaciations, the Lines: 78 to 82 16 Missouri River had flowed northeastward into Hudson Bay, probably cutting 17 across northern Montana and entering Canada in southeastern Saskatchewan. ——— 18 Its northern tributaries, including the Yellowstone, the Little Missouri, and the 0.0pt PgVar ——— 19 now isolated Souris, were part of this same general river system, as perhaps Normal Page 20 also were the Knife and the Cannonball rivers. Then, about 2 million years PgEnds: T X 21 ago, the first of a series of continent-wide glaciations developed across most E 22 of northern North America, with roughly the northeastern fifth of the Great 23 Plains becoming repeatedly covered with sheets of ice hundreds of feet thick. At [4], (4) 24 maximum, the glaciers’ southernmost limits in the Great Plains extended from 25 the general vicinity of the Rocky Mountain foothills in present-day northwest- 26 ern Montana eastward and southward along a line approximating the present 27 southeastward-flowing course of the Missouri River, to eastern Nebraska and 28 northeastern . 29 With the glaciers came a host of cold-adapted mammals, including masto- 30 dons, mammoths, bison, caribou, muskoxen, wolves, tiger-sized cats, bears, and 31 many other cold-adapted species (Schultz 1934). Along the glaciers’ southwest- 32 ern edges in what is now the Dakotas, ridges of rock debris produced terminal 33 moraines that are collectively called the Coteau du Missouri. Irregularities left 34 by stagnant ice produced pockmarked terrains in northeastern Montana and 35 the Dakotas that eventually would become the famous“duck factory”or“prairie 36 pothole” region. The Missouri River began to form its present, generally south- 37 ward flowing course, and rivers like the Platte and Niobrara also shifted their 38 courses as they were variously influenced by glacial moraines and meltwater 39 effects. The current channel of the Missouri closely follows the southwestern 40 limits of the last (Wisconsinian) glaciation. Highly sculptured badlands in the

4 the western shortgrass prairie

Kim—UofNPress/Page4//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 western Dakotas were also produced, mostly by water erosion from glacier-fed 2 rivers that now sometimes are barely more than seasonal trickles. 3 The plant and animal life present during this broad swatch of NorthAmerican 4 history was probably an ever-changing tapestry, as climate patterns ebbed and 5 flowed and as the land itself was reshaped. It is clear that the general pattern of 6 climatic change in the Great Plains, as elsewhere in North America, has been one 7 of general cooling and drying, at least since the end of the Age of Dinosaurs (the 8 Cretaceous era) some 65 million years ago. As mentioned, the tropical forests 9 that once covered the great interior lowlands of North America disappeared 10 and were very gradually replaced by more - and cold-tolerant forms. 11 Thus tropical evergreen trees were replaced by seasonably deciduous trees, de- 12 ciduous trees by shrubs, and shrubs by herbs (broad-leaved herbaceous plants 13 and grasses). Grasses, the plant type best adapted to short growing seasons, [5], (5) 14 recurrent fires, and grazing by large animals, were destined to become the dom- 15 inant plant type of the Plains. Somewhere between the middle Miocene epoch Lines: 82 to 88 16 (about 15 million years ago) and historic times, the Great Plains were converted 17 from a semiopen forest, with only scattered grassy areas, to a completely open ——— 18 grassland, with forests and woodlands limited to watercourses and other areas 0.0pt PgVar ——— 19 protected from fire and having access to surface or subsurface moisture (Axelrod Normal Page 20 1985). PgEnds: T X 21 Plants living in northwestern Mexico, a region of long-term deserts and E 22 semideserts associated with the permanent high-pressure zones of these lati- 23 tudes, probably provided a reservoir of plants and animals pre-adapted to an [5], (5) 24 arid-zone pattern of existence. From this general region, especially the present- 25 day Chihuahuan Desert, various grasses and other ecologically similar plants 26 presumably moved north to occupy the increasingly significant rain-shadow 27 effect of the Rocky Mountains, where most of the clouds carrying moisture 28 from the Pacific Ocean were nearly squeezed dry as they passed eastward over 29 the high western cordilleras. 30 The lowly but pervasive buffalo grass, which now extends from the south- 31 ern tip of the Chihuahuan Desert north across the drier plains almost to the 32 Canadian border, is one of the many arid-adapted grasses that probably had 33 its origins here, and it has certainly been a foraging mainstay for such grazing 34 mammals as bison and prairie dogs for uncountable millennia. The Mexican 35 prairie dog is a southern representative of the keystone prairie dog assemblage of 36 five grassland-dependent species and likewise is a part of this same longstanding 37 semidesert community. It also is perhaps a Mexican relict of an original arid 38 grassland distribution pattern of ancestral prairie dogs. It still survives in a 39 southern corner of the Chihuahuan Desert, although it is now an endangered 40 species. The coyote may have originated in the Mexican plateau too; in the

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Kim—UofNPress/Page5//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 Aztec culture, Heuheucoyotl was a trickster deity and Coyolxauhqui was the 2 moon goddess. Our English name for the coyote is thus derived from the Aztec 3 coyotl, a fitting tribute both to its cunning nature and to its love for singing on 4 moonlit nights. Likewise the kit fox, a close kin to the more northerly swift fox, 5 has a semitropical desert grassland orientation, as do several jackrabbit species, 6 such as the white-sided jackrabbit. The adjacent dry mountain slopes of the 7 Sierra Madre Occidental and Oriental, where scrub oaks, mesquite, and similar 8 arid-adapted woody plants still occur, probably provided various shrubs and 9 deciduous trees to the mix of plant species that were slowly inching their way 10 northward over the western plains. 11 The climate of the western plains gradually settled into a pattern character- 12 ized by dry, cold winters, with much of the cold-season falling as [6], (6) 13 snow. Most of the annual precipitation occurred during the spring and sum- 14 mer growing season, often in the form of violent thunderstorms that occurred 15 as moisture-laden warm air from the Gulf of Mexico met cooler and drier air Lines: 88 to 102 16 currents coming southeast out of the northern interior. Increasingly short sum- 17 ——— mers occurred northwardly, and increasing amounts of annual sunshine and 18 13.0pt PgVar associated evaporation occurred toward the south. However, periodic droughts ——— 19 and recurrent, lightning-caused prairie fires added to the complexities of life Normal Page 20 for organisms adapting to these demanding conditions for survival. The grasses PgEnds: T X 21 E and other plants of the developing shortgrass and mixed-grass prairies were 22 ones that, like the desert and desert grassland plants of the Southwest, were 23 [6], (6) mainly warm-season perennial species, whose photosynthetic processes oper- 24 ate well under conditions of high summer temperatures and extended periods 25 26 of relatively little moisture, sometimes lasting for several years. Among these 27 species are such important southern and western shortgrass and mixed-grass 28 prairie genera as Andropogon, Buchloë, Bouteloua, Chloris, Paspalum, Setaria, 29 Schizachyrium, Sorghastrum, and Sporobolus. Plants that evolved to form the 30 tallgrass prairies at the northeastern, moister edges of the Great Plains have 31 a higher proportion of cool-season grass genera. These include such typical 32 prairie grasses as Agropyron, Bromus, Elymus, Festuca, Hordeum, Poa, and Stipa 33 (Sims 1988). At the northernmost edge of the Great Plains, Festuca grasses 34 became especially important in the transition zone toward the forests of the 35 Rockies and the Canadian boreal forests. These tall, productive grasses are well 36 adapted to cooler summer temperatures and greater precipitation than are most 37 mixed-grass prairie grasses. As a result, bison of the Canadian plains histori- 38 cally moved west toward the Front Range of the Rocky Mountains, rather than 39 migrating great distances southward, to obtain their fall and winter sustenance 40 from this nutritious forage source.

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Kim—UofNPress/Page6//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 the land today 2 3 The present climate of the western plains may be summarized relative to its 4 geographically variable precipitation.At the western edge of the plains an annual 5 precipitation of less than 16 inches is typical. The mixed-grass prairies merge 6 eastwardly with the tallgrass prairies very roughly at the isohyet of about 28 7 inches of annual precipitation. 8 Various ways exist to represent the geography, climate, and associated bi- 9 ological communities of the Great Plains visually. One current and widely 10 adopted method was first proposed by Robert Bailey. It recognized twenty-five 11 separate ecological-geographic provinces in the United States south of Canada. 12 Bailey’s boundaries were refined by The Nature Conservancy (1999) into fifty- 13 two ecoregions within the same forty-eight-state boundaries, of which five fall [7], (7) 14 within the previously designated geographic limits of the western plains (Map 15 1). Of these five the Black Hills are not geologically a part of this region, leaving Lines: 102 to 120 16 the Northern Great Plains , the Central Mixed-grass Prairie, the Central 17 Shortgrass Prairie, and the Southern Shortgrass Prairie as constituent parts of ——— 18 the western plains. Some of the limits of these ecoregions have since undergone 0.0pt PgVar ——— 19 further minor refining, but the collective boundaries of these five ecoregions Normal Page 20 make for a convenient visualization of the entire western plains biological do- PgEnds: T X 21 main and will be used later in the book as a collective biogeographical definition E 22 of this region. 23 More recently a group of avian biologists formed the North American Bird [7], (7) 24 Conservation Initiative Committee (nabcic), which operates through region- 25 ally based, biologically driven, and landscape-oriented partnerships. A team 26 from that group assembled a map of proposed Bird Conservation Regions 27 (North American Bird Conservation Initiative Committee 2000). This map di- 28 vides the Northern Great Plains Steppe into a more northerly“Prairie Potholes” 29 component, which extends north into central Alberta and east into the tallgrass 30 prairies of the Dakotas, Minnesota, and Iowa, and a more southerly and drier 31 “Badlands and Prairies”segment. The other western plains elements recognized 32 in the map are the southwestern Shortgrass Prairie region and the southeastern 33 Central Mixed-grass Prairie segment. Three of these Bird Conservation Regions 34 are mapped here (Map 2); the map excludes the Prairie Potholes segment, as it 35 mostly falls outside the recognized geographic limits of the western plains. 36 The last map reproduced here includes the entire Great Plains and part of the 37 Central Lowlands of North America (Map 3). It is largely based on a National 38 Wildlife Federation historical vegetation-based map (Bachard 2001), which is 39 geographically limited to the native perennial grasslands (tallgrass, mixed-grass, 40 and shortgrass prairies) of interior North America. Additionally, the Nebraska

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Kim—UofNPress/Page7//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 Sandhills are designated separately in Map 3; they are botanically and zoolog- 2 ically quite recognizable, and their separate inclusion provides a convenient 3 geographic separation between the biologically distinguishable northern and 4 southern segments of the mixed-grass prairies. 5 It is one thing to look at a map and try to visualize the kinds of landscapes 6 represented therein, to say nothing of the diversity of plants and animals that 7 might be implied in a simple map label such as“shortgrass prairie.”For example, 8 the native prairies of Buffalo Gap National Grassland in the southwestern South 9 Dakota badlands region are known to support 230 species of birds, 58 mammals, 10 19 reptiles, and 8 amphibians as well as 92 grasses, 160 broad-leaved herbs (or 11 forbs), 22 shrubs, and 11 trees. Beyond these are uncountable numbers of insects, 12 other invertebrates, and soil microorganisms. Most of this very land was once 13 ranched or farmed for only a few decades, from when it was first settled in the [8], (8) 14 later 1800s until the years of the 1930s, when bankruptcies became as 15 common as summer dust storms. Plowing up such fragile lands to raise Lines: 120 to 146 16 or corn for a few decades, often until the topsoil blows away and the land is 17 abandoned, is like throwing away a treasure trove of potential biological riches ——— 18 to raise a single species of grass that needs so much tilling, water, herbicides, * 218.58002pt PgVar ——— 19 and pesticides to survive that scarcely anything else of value can survive there. Normal Page 20 * PgEnds: PageBreak 21 One’s native land is the most important thing on earth. Above all it is made 22 holy by the ancestors, who pass it on. – Louis Tiel 23 [8], (8) 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

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Kim—UofNPress/Page8//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 2 3 ALBERTA 4 SASKATCHEWAN 5 6 7 8 9 NORTH 10 DAKOTA 11 MONTANA Northern Great 12 Plains Steppe 13 [9], (9) 14 Black Hills 15 SOUTH DAKOTA Lines: 146 to 170 16 WYOMING 17 ——— 18 8.992pt PgVar ——— 19 NEBRASKA Normal Page 20 PgEnds: T X 21 E 22 Central Central 23 Shortgrass Mixed- KANSAS [9], (9) 24 Prairie grass 25 Prairie 26 27 28 Southern OKLAHOMA 29 Shortgrass 30 Prairie NEW MEXICO 31 32 33 34 TEXAS 35 36 37 0 Miles 500 38 39 Ecoregions of the Great Plains. After the Nature Conservancy (1999). 40

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Kim—UofNPress/Page9//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 2 3 4 5 6 7 8 9 MONTANA 10 NORTH 11 DAKOTA 12 Badlands & Prairies 13 [10], (10) 14 SOUTH 15 DAKOTA Lines: 170 to 170 16 17 WYOMING ——— 18 -1.342pt PgVar ——— 19 Normal Page 20 NEBRASKA PgEnds: T X 21 Central E 22 COLORADO Mixed-grass 23 Prairie [10], (10) 24 Shortgrass KANSAS 25 Prairie 26 27 28 OKLAHOMA

29 NEW MEXICO 30 31 32 33 34 35 TEXAS 36 37 0 Miles 500 38 39 Bird Conservation Regions of the High Plains. After the North American Bird Conser- 40 vation Initiative Committee (2000).

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Kim—UofNPress/Page10//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard 1 ALBERTA MANITOBA 2 SASKATCHEWAN 3 4 5 6 7 8 9 Mixed-grass Prairie MINNESOTA 10 NORTH MONTANA DAKOTA 11 [Last Page] 12 [11], (11) 13 SOUTH 14 DAKOTA 15 WYOMING Lines: 170 to 174 16 Sandhills IOWA 17 Prairie ——— 18 -2.008pt PgVar ——— 19 NEBRASKA Normal Page 20 COLORADO Mixed- PgEnds: TEX 21 grass KANSAS 22 Prairie MISSOURI 23 [11], (11) 24 Shortgrass 25 Prairie OKLAHOMA 26 NEW MEXICO 27 28 TEXAS 29 30 31 32 33 34 35 Coastal Tallgrass Prairie 36 Miles 37 0 500 38 39 Biological regions of the Great Plains. Mainly after Bachand (2001) but with the Sandhills 40 Prairie additionally designated.

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Kim—UofNPress/Page11//Prairie Dog Empire: A Saga of the Shortgrass Prairie / Paul A. Johnsgard