Freshwater Fishes of North America

Home , Acanthopterygii, Acipenseriformes, Actinopteri, Amblyopsidae, Amiiformes, Atheriniformes

© 2019 The Johns Hopkins University Press UNCORRECTED PROOF Do not quote for publication until verified with finished book All rights reserved. No portion of this may be reproduced or distributed without permission. NOT FOR SALE OR DISTRIBUTION

349-82339_Warren_ch00_1P.indd 1 10/24/19 10:32 PM Freshwater Fishes of North Amer i ca VOLUME 2 Characidae to Poeciliidae Edited by Melvin L. Warren, Jr., and Brooks M. Burr With Anthony A. Echelle, Bernard R. Kuhajda, and Stephen T. Ross Illustrated by Joseph R. Tomelleri

Johns Hopkins University Press BALTIMORE

349-82339_Warren_ch00_1P.indd 3 10/24/19 10:32 PM This page intentionally left blank Freshwater Fishes of North Amer i ca VOLUME 2 Characidae to Poeciliidae Edited by Melvin L. Warren, Jr., and Brooks M. Burr With Anthony A. Echelle, Bernard R. Kuhajda, and Stephen T. Ross Illustrated by Joseph R. Tomelleri

Johns Hopkins University Press BALTIMORE

Johns Hopkins University Press 2715 North Charles Street Baltimore, Mary land 21218-4363 www . press . jhu . edu

Library of Congress Cataloging- in- Publication Data

Freshwater ﬁshes of North Amer i ca / edited by Melvin L. Warren, Jr., and Brooks M. Burr ; illustrated by Joseph R. Tomelleri. volumes cm Includes bibliographical references and index. ISBN-13: 978-1-4214-1201-6 (hardcover : alk. paper) ISBN-13: 978-1-4214-1202-3 (electronic) ISBN-10: 1-4214-1201-2 (hardcover : alk. paper) ISBN-10: 1-4214-1202-0 (electronic) 1. Freshwater ﬁshes—North Amer i ca. I. Warren, Melvin L., Jr., editor of compilation. II. Burr, Brooks M., editor of compilation. QL625.F74 2014 597.176— dc23 2013015264 ISBN-13: 978-1-4214-3512-1 (hardcover : alk. paper) ISBN-13: 978-1-4214-3513-8 (electronic) ISBN-10: 1-4214-3512-8 (hardcover : alk. paper) ISBN-10: 1-4214-3513-6 (electronic)

A cata log record for this book is available from the British Library.

Special discounts are available for bulk purchases of this book. For more information, please contact Special Sales at 410-516 -6936 or specialsales@press . jhu . edu.

Johns Hopkins University Press uses environmentally friendly book materials, including recycled text paper that is composed of at least 30 percent post- consumer waste, whenever pos si ble.

349-82339_Warren_ch00_1P.indd 4 10/24/19 10:32 PM Robert C. Cashner (1941–2018) Friend, teacher, mentor, leader, and stellar freshwater ichthyologist. We miss you, Bob.

349-82339_Warren_ch00_1P.indd 5 10/24/19 10:32 PM This page intentionally left blank Contents

List of Contributors ix Preface xi Melvin L. Warren, Jr., and Brooks M. Burr Acknowl edgments xix

Chapter 14 Characidae: Characins 1 Norman Mercado- Silva, Claudia P. Ornelas- García, Juan Jacobo Schmitter- Soto, Nicholas J. Gidmark, and Andrew M. Simons

Chapter 15 Ictaluridae: North American Catﬁshes 23 Brooks M. Burr, Melvin L. Warren, Jr., and Micah G. Bennett

Chapter 16 Ariidae: Sea Catﬁshes 101 Matthew R. Thomas and Ricardo Betancur-R.

Chapter 17 Heptapteridae: Seven- ﬁnned Catﬁshes 123 Matthew R. Thomas and Mark Henry Sabaj

Chapter 18 Osmeridae: Smelts 149 Robert L. Hopkins II and Melvin L. Warren, Jr.

Chapter 19 Esociformes: Esocidae, Pikes, and Umbridae (Mudminnows) 193 Frank H. McCormick, Terry Grande, Cheryl Theile, Melvin L. Warren, Jr., J. Andrés López, Mark V. H. Wilson, Roger A. Tabor, Julian D. Olden, and Lauren M. Kuehne

Chapter 20 Percopsidae: Trout- perches 261 Brooks M. Burr and Melvin L. Warren, Jr.

Chapter 21 Amblyopsidae: Caveﬁshes 281 Ginny L. Adams, Brooks M. Burr, and Melvin L. Warren, Jr.

Chapter 22 Aphredoderidae: Pirate Perches 322 Brooks M. Burr and Melvin L. Warren, Jr.

349-82339_Warren_ch00_1P.indd 7 10/24/19 10:32 PM viii COnTEnTs

Chapter 23 Gadidae (Gadinae and Lotinae): Cods and Cuskfishes 340 Peter B. Berendzen

Chapter 24 Mugilidae: Mullets 367 Steven L. Powers

Chapter 25 Atherinopsidae: New World Silversides 384 Kyle R. Piller and Clyde D. Barbour

Chapter 26 Beloniformes: Belonidae (Needlefishes) and Hemiramphidae (Halfbeaks) 449 Bruce B. Collette and Stephen J. Walsh

Chapter 27 Rivulidae: New World Rivulines 463 Ann M. Uzee O’Connell, Martin T. O’Connell, and Anthony A. Echelle

Chapter 28 Profundulidae: Middle American Killifishes 493 Mollie F. Cashner and Anthony A. Echelle

Chapter 29 Goodeidae: Goodeids 503 Shane A. Webb

Chapter 30 Fundulidae: Topminnows 549 Robert C. Cashner, Jacob Schaefer, Melvin L. Warren, Jr., Anthony A. Echelle, Fernando Galvez, and Michael J. Ghedotti

Chapter 31 Cyprinodontidae: Pupfishes 609 Anthony A. Echelle and Alice F. Echelle

Chapter 32 Poeciliidae: Livebearers 674 Michael J. Ghedotti, Matthew P. Davis, and Anthony A. Echelle

Lit er a ture Cited 739 Index of Scientific Names 889 General Index 898

349-82339_Warren_ch00_1P.indd 8 10/24/19 10:32 PM Contributors

Ginny L. Adams Michael J. Ghedotti Steven L. Powers University of Central Arkansas Regis University Roanoke College Clyde D. Barbour (retired) Nicholas J. Gidmark Mark Henry Sabaj Wright University Knox College The Academy of Natu ral Sciences Micah G. Bennett Terry Grande of Drexel University U.S. Environmental Protection Agency Loyola University Chicago Jacob Schaefer Peter B. Berendzen Robert L. Hopkins II University of Southern Mississippi University of Northern Iowa University of Rio Grande Juan Jacobo Schmitter- Soto Ricardo Betancur-R. Lauren M. Kuehne El Colegio de la Frontera Sur University of Puerto Rico University of Washington Andrew M. Simons Brooks M. Burr J. Andrés López University of Minnesota Southern Illinois University University of Alaska Roger A. Tabor Mollie F. Cashner Frank H. McCormick U.S. Fish and Wildlife Ser vice Austin Peay State University USDA Forest Ser vice Cheryl Theile Robert C. Cashner (deceased) Norman Mercado- Silva Loyola University Chicago University of New Orleans Universidad Autónoma del Estado Matthew R. Thomas Bruce B. Collette de Morelos Kentucky Department of Fish National Museum of Natu ral History Ann M. Uzee O’Connell and Wildlife Resources Matthew P. Davis University of New Orleans Stephen J. Walsh St. Cloud State University Martin T. O’Connell U.S. Geological Survey Alice F. Echelle University of New Orleans Melvin L. Warren, Jr. Oklahoma State University Julian D. Olden USDA Forest Ser vice Anthony A. Echelle University of Washington Shane A. Webb Oklahoma State University Claudia P. Ornelas- García University of North Georgia Fernando Galvez Universidad Nacional Autónoma de Mark V. H. Wilson (emeritus) Louisiana State University México Tercer Circuito Exterior University of Alberta Kyle R. Piller Southeastern Louisiana University

349-82339_Warren_ch00_1P.indd 9 10/24/19 10:32 PM This page intentionally left blank Preface

The North American freshwater fish fauna comprises knowledge of the North American freshwater fish fauna >1,300 native species in about 50 families. It is the most theretofore unknown. Shortly thereafter a physician from thoroughly studied and largest temperate fish fauna Forsythe, Missouri, combined his hobby of scuba diving (Miller et al. 2005; Page & Burr 2011) in the world. In and snorkeling with photography and revealed— new even comparison, an analy sis and compendium of Euro pean to specialists— the incredible colors of the North American freshwater fishes included 546 native species in about 24 native fish fauna, especially in their brightest breeding con- families (Kottelat & Freyhof 2007); Eu rope is about one- dition, as well as some of their unique and fascinating natu- third the land area of North Amer i ca. Australia has nearly ral histories. William N. Roston eventually traveled the 300 freshwater fishes in 35 families (Allen 1989; Allen continent looking for clear water and fish to photo graph in et al. 2002) in a land area about that of the United States their natu ral environment (never in aquaria). A number of (minus Alaska). This number includes many marine spe- his photo graphs are used in this volume. cies that enter fresh water, and highly unusual freshwater From the 1970s through the 1990s numerous books fish lineages occur there (e.g., Salamanderfish, Lepidogal- focused on fish faunas of individual states (e.g., Alabama, axias salamandroides; Australian Lungfish, Neoceratodus Arkansas, California, Idaho, Illinois, Kansas, Mississippi, forsteri; and Nurseryfish, Kurtus gulliveri). The only other Missouri, New Mexico, Ohio, Tennessee, Virginia, Wash- temperate fish fauna that could rival North Amer i ca is ington, West Virginia, and Wisconsin) as well as Canada, Asia (Lévêque et al. 2008), but reliable information on making even more detail on fishes available to the public. this vast area and its fishes remains poorly understood by These works allowed for the first complete identification scientists in the New World. Estimates for the country of guide to all freshwater fishes in the United States and China range from 920 to 1,010 native species (Kang et al. Canada (Page & Burr 1991, revised 2011). Nevertheless, it 2014; M. Kottelat pers. comm.). Unsurprisingly, as for was not until the Freshwater Fishes of Mexico (Miller et al. many plant and animal groups, the tropical regions of the 2005, 2009) was published that it was possi ble for us to world harbor freshwater fish faunas several times larger consider editing this three- volume work on the natu ral than those of temperate regions (Lundberg et al. 2000; history, ecol ogy, and conservation of North American Agostinho et al. 2005; Berra 2007). freshwater fishes. We are indebted to a large community In the mid-1970s, knowledge of North American fresh- of ichthyologists, fisheries biologists, and other workers in water fishes was confined to a few specialists, but even so related fields (e.g., ge ne tics, physiology, behav ior, ecology) for many species (and families) little was available on natu- who have investigated the details of the lives of fishes in ral history or ecol ogy. In 1980 a landmark volume was pub- such a way that much technical information can now be lished that used spot distribution maps to illustrate the synthesized in one place and again made available to the ranges of all freshwater fish species in the United States public and other specialists. and Canada (Lee et al. 1980 et seq.). That volume made Even though our overarching goal was to synthesize as available to the lay public as well as specialists a level of much information as pos si ble on North American freshwater

349-82339_Warren_ch00_1P.indd 11 10/24/19 10:32 PM xii PREFACE

fishes, the job of gathering information is far from complete. systems (e.g., Striped Mullet, Mugil cephalus, Powers, this In editing this work and writing synthesis chapters of our volume). Admittedly, some degree of arbitrariness was un- own, we were struck at once by the incredible natu ral history avoidable in the inclusion or exclusion of marine species and taxonomic diversity among our native freshwater fishes, in a work about freshwater fishes, but the arbitrariness re- but also by the large and critical information gaps that re- flects a biologically real gray area among fishes at the in- main. Unfortunately, for many species (and nearly entire terface of freshwater and saltwater systems (e.g., coastal families), the syntheses presented here are (or are close to be- Largemouth Bass, Micropterus salmoides; Gulf Pipefish, ing) obituaries (e.g., Miller et al. 1989; Minckley & Deacon Syngnathus scovelli; Atlantic Needlefish, Strongylura ma- 1991; Warren et al. 2000; Brouder & Scheurer 2007; Jelks rina, Collette & Walsh, this volume). et al. 2008; Burkhead 2012). For many species and groups, Each taxonomic chapter focuses on a family or in some the biological information needed to help recover them, to cases two families of North American freshwater fishes slow population declines, or to prevent extinction is simply with emphasis on the natu ral history, biology, evolution, unavailable. That said, the most critical component of con- and conservation of each genus in the family. Because of serving North American freshwater fishes is the prevention the timing of preparation of the volumes, the sequence of of habitat loss and degradation by humans, not the lack of bi- the families generally follows the arrangement of Page ological information. Fishes in this fauna have an incredible et al. (2013), Nelson (2006), and Nelson et al. (2016), but tenacity for life, whether we completely understand their we acknowledge the phylogeny of Ray- finned Fishes and biology or not, but we as coinhabitants of the North Ameri- Spiny- rayed Fishes by Near et al. (2012b, 2013) and the ex- can continent need to provide them the opportunity to en- pansion of that work by Betancur- R. et al. (2013ab, 2017) dure. We hope that this work helps stem the high rates of as the most comprehensive and defensible to date and pre- population decline and extinction being experienced across sent these sequences herein for North American fishes we the North American fish fauna. We also hope this work stim- cover ( Tables P.1 and P.2). In Volume 1, taxonomic chap- ulates a whole new generation of ichthyologists and fisheries ters covered the Lampreys (Petromyzontidae) through the researchers to further expand our knowledge and apprecia- Suckers (Catostomidae) with one exception. Taxonomic tion of the natu ral history, ecol ogy, and conservation of the chapters in Volume 2 cover the Characins (Characidae) great freshwater fish fauna of North Amer i ca. through the Livebearers (Poecilidae) with one exception, and in Volume 3, the Sticklebacks through the American AREA AND BREADTH OF COVERAGE Soles (Achiriidae). Because of extenuating circumstances, the chapters on Herrings (Clupeidae; originally slated for The area of coverage encompasses fishes in fresh waters Volume 1) and Trouts and Salmons (Salmonidae; originally of North Amer i ca, including Canada, the coterminous slated for Volume 2) will appear in Volume 3. United States, and Mexico, south generally to the Isthmus of Tehuantepec. For some families, authors extended the METHODS southern boundary to include fishes of the Yucatan Penin- sula region. Within the covered area, all native North Sources. We encouraged authors to use only peer- American fishes, emphasizing the level of genus, are in- reviewed publications in this work, but in many cases, included that primarily inhabit and reproduce in fresh formation was only available in unpublished dissertations, water and that primarily inhabit marine or estuarine sys- theses, or even reports. We clearly indicate those unpub- tems but are frequent or even permanent components of lished sources as such in the lit er a ture cited. some freshwater fish assemblages. Some primarily marine Scientific and common names. For scientific and com- fishes (across 11 families) are included because they are mon names of taxa (e.g., species, genera, families, orders), naturally established and reproduce in fresh waters (e.g., we used Nelson (2006), Page et al. (2013), Nelson et al. Atlantic Needlefish, Strongylura marina, Collette & Walsh, (2016), and occasionally FishBase (Froese & Pauly 2019) this volume). Others are included because young and as guides. Our order of pre sen ta tion of families generally adults penetrate deeply into freshwater systems and re- follows that of Nelson (2006), Page et al. (2013), and produce or are suspected of reproducing in fresh waters Nelson et al. (2016), but as noted previously, Near et al. (e.g., Hogchoker, Trinectes maculatus), and still others sim- (2012b, 2013) presented a phylogeny of Ray- finned Fishes ply occur with such high frequency in freshwater habitats (Actinopterygii) and Spiny- rayed Fishes (Acanthomorpha) that they likely are impor tant functionally in those eco- based on 9–10 nuclear genes and 579 fish species. Using

349-82339_Warren_ch00_1P.indd 12 10/24/19 10:32 PM Table P.1. Phyloge ne tic sequence, clade names, orders, and family names of Ray- finned Fishes (Actinopterygii) and Spiny- rayed Fishes (Acanthopterygii) represented in North American fresh waters. The sequence follows the phyloge ne tic trees recovered from analy sis of 9–10 nuclear genes and 579 fish species (modified from Near et al. 2012b, 2013). The designation at the ordinal level of incerti ordis indicates the broader relationships of the family are to date undefined. Clade Name Order and Family (common name) Actinopterygii Actinopteri Acipenseriformes Acipenseridae (Sturgeons) Polyodontidae (Paddlefishes) Neopterygii Holostei Amiiformes Amiidae (Bowfins) Lepisosteiformes Lepisosteidae (Gars) Teleostei Elopomorpha Anguilliformes Anguillidae (Freshwater Eels) Osteoglossocephala Osteoglossomorpha Hiodontiformes Hiodontidae (Mooneyes) Clupeocephala Otocephala Clupeiformes Clupeidae (Herrings) Engraulidae (Anchovies) Ostariophysi Otophysi Cypriniformes Catostomidae (Suckers) Cyprinidae (Carps and Minnows) Characiformes Characidae (Characins) Siluriformes Ariidae (Sea Catfishes) Heptapteridae (Seven- finned Catfishes) Ictaluridae (North American Catfishes) Euteleostei Salmoniformes Salmonidae (Trouts and Salmons) Esociformes Esocidae (Pikes and Mudminnows) Osmeriformes Osmeridae (Smelts) Neoteleostei Eurypterygii Ctenosquamata Acanthomorpha Percopsiformes Amblyopsidae (Cavefishes) Aphredoderidae (Pirate Perches) Percopsidae (Trout- Perches) Gadiformes Gadidae (Cods) (continued)

xiii

349-82339_Warren_ch00_1P.indd 13 10/24/19 10:32 PM xiv PREFACE

Table P.1, continued Clade Name Order and Family (common name) Acanthopterygii Percomorpha Syngnathiformes Syngnathidae (Pipefishes and Sea horses) Gobiiformes Gobiidae (Gobies) Eleotridae (Sleepers) Synbranchiformes Synbranchidae (Swamp Eels) Pleuronectiformes Achiridae (American Soles) Paralichthyidae (Sand Flounders) Pleuronectidae (Righteye Flounders) Ovalentaria Atheriniformes Atherinopsidae (New World Silversides) Beloniformes Belonidae (Needlefishes) Hemiramphidae (Halfbeaks) Cyprinodontiformes Rivulidae (New World Rivulines) Goodeidae (Goodeids) Profundulidae ( Middle American Killifishes) Cyprinodontidae (Pupfishes) Fundulidae (Topminnows) Poeciliidae (Livebearers) incerti ordinis Cichlidae (Cichlids and Tilapias) incerti ordinis Mugilidae (Mullets) Embiotocidae (Surfperches) incerti ordinis Gobiesocidae (Clingfishes) Unnamed clade Centrarchiformes Centrarchidae (Sunfishes) Perciformes Percidae (Perches) Cottidae (Sculpins) Gasterosteidae (Sticklebacks) Unnamed clade incerti ordinis Moronidae (Temperate Basses) Unnamed clade incerti ordinis Sciaenidae (Drums)

and expanding and updating the Near et al. (2013) data, the chapters on first mention of the species and thereafter Betancur- R. et al. (2013ab, 2017) analyzed relationships of use either or both. Authors were free to deviate from nearly 2,000 bony fish taxa using multi- locus trees. The these primary sources for common and scientific names results clearly indicate convergence on a well-resolved for newly described species and higher taxa or because of phylogeny for most fishes. We accept those works as defin- di«ering or new systematic evidence (or taxonomic opin- itive ( Tables P.1 and P.2) but did not follow that phyloge- ion) as well as for clarity. We capitalized the common ne tic sequence because of timing of volume preparation. names, if available, of all species, families, orders, and Authors give the complete scientific and common name in higher taxonomic categories (e.g., Ray- finned Fishes for

349-82339_Warren_ch00_1P.indd 14 10/24/19 10:32 PM Table P.2. Phyloge ne tic classification of Ray- finned Fishes (Actinopterygii) represented in North American fresh waters. The sequence follows the phyloge ne tic tree recovered from analy sis of multiple locus DNA sequence data for nearly 2,000 fish taxa (Betancur- R. et al. 2013ab, 2017). The designation at the ordinal level of incerti ordinis indicates the broader relationships of the family are to date undefined. Class Subclass Infraclass Megacohort Superorder Order Family (common name) Actinopteri Chondrostei Acipenseriformes Acipenseridae (Sturgeons) Neopterygii Polyodontidae (Paddlefishes) Holostei Amiiformes Amiidae (Bowfins) Lepisosteiformes Lepisosteidae (Gars) Teleostei Elopocephali Anguilliformes Anguillidae (Freshwater Eels) Osteoglossocephalai Hiodontiformes Hiodontidae (Mooneyes) Clupeiformes Clupeidae (Herrings) Engraulidae (Anchovies) Cypriniphysae Cypriniformes Catostomidae (Suckers) Cyprinidae (Carps and Minnows) Characiformes Characidae (Characins) Siluriphysae Siluriformes Ariidae (Sea Catfishes) Heptapteridae (Seven- finned Catfishes) Ictaluridae (North American Catfishes) Salmoniformes Salmonidae (Trouts and Salmons) Esociformes Esocidae (Pikes) Umbridae (Mudminnows) Osmeriformes Osmeridae (Smelts) Percopsiformes Amblyopsidae (Cavefishes) Aphredoderidae (Pirate Perches) Percopsidae (Trout- perches) Gadiformes Gadidae (Cods and Cuskfishes) Gobiiformes Eleotridae (Sleepers) Gobiidae (Gobies) Syngnathiformes Syngnathidae (Pipefishes and Sea horses) Synbranchiformes Synbranchidae (Swamp Eels) Pleuronectiformes Achiridae (American Soles) Paralichthyidae (Sand Flounders) Pleuronectidae (Righteye Flounders) incerti ordinis Embiotocidae (Surfperches) Atherinomorphae Atheriniformes Atherinopsidae (New World Silversides) Cichlomorphae Cichliformes Cichlidae (Cichlids and Tilapias) Atherinomorphae Beloniformes Belonidae (Needlefishes) Hemiramphidae (Halfbeaks)

Atherinomorphae Cyprinodontiformes Cyprinodontidae (Pupfishes) Fundulidae (Topminnows) Poeciliidae (Livebearers) Goodeidae (Goodeids) Profundulidae ( Middle American Killifishes) “Rivulidae” (New World Rivulines) Mugilomorphae Mugiliformes Mugilidae (Mullets) Blenniimorphae Gobiesociformes Gobiesocidae (Clingfishes) incerti ordinis Moronidae (Temperate Basses) incerti ordinis Sciaendae (Drums) Centrarchiformes Centrarchidae (Sunfishes) Elassomatidae (Pygmy Sunfishes) Perciformes Percidae (Perches) Gasterosteidae (Sticklebacks) Cottidae (Sculpins)

349-82339_Warren_ch00_1P.indd 15 10/24/19 10:32 PM xvi PREFACE

Actinopterygii, Catfishes for Siluriformes, North Ameri- 1992; Ross 2013; Ross & Matthews 2014). Likewise, con- can Catfishes for Ictaluridae, Channel Catfish for Ictalurus tributors did not include tools or aids in identification of punctatus) (see Nelson et al. 2002). We encouraged au- species (e.g., illustrated keys) because identification is most thors to use the common name in lieu of scientific names often a species- level exercise and it is so well covered in after first mention because common names are descrip- numerous state and regional fish books, including Canada tive and colorful and are increasingly more stable through (Scott & Crossman 1973) and Mexico (Miller et al. 2005, time than scientific names. We did not capitalize collo- 2009; see also lit er a ture guide sections in each taxonomic quial, nonstandard, or semi- technical, but informal, com- chapter) as well as in a field guide for North American mon names for groups of fish (e.g., bass, blackbass, brook freshwater fishes north of Mexico (Page & Burr 2011). An- lamprey, bu«alo, bullhead, carp, carpsucker, chub, crap- other impor tant, but large topic not covered in detail is the pie, dace, darter, jumprock, killifish, madtom, minnow, area of fishing statistics, which again is deserving of a sep- molly, mosquitofish, pickerel, pikeminnow, piranha, red- arate synthesis (see commercial importance sections for horse, salmon, shad, shiner, splitfin, trout). some details). We describe the content of major sections Fossil taxa. We indicated fossil taxa by a dagger (†) placed and subsections in each chapter as follows: before the genus name. In general, we followed Walker Chapter introduction. In an initial section, contributing et al. (2012) for designation of geologic time (period, epoch, authors introduce the family to the reader by relating the age) in millions of years ago (mya), but the original refer- scientific and common names of the fishes, highlighting ences should be consulted to determine how the geological some specialized or unusual features of the group, and for formations were dated or how fossil dates were estimated. those families with a large number of marine species or Length abbreviations and museum acronyms. We abbrevi- those not wholly endemic to fresh waters of North Amer- ated standard length (SL), total length (TL), and fork i ca, placing the focal taxa in context of the diversity and length (FL) throughout. Museum acronyms followed geographic distribution of the entire family. Sabaj Pérez (2016). Diversity and distribution. Contributing authors summa- Distributional maps. We provided to authors shaded maps rize the general diversity of the focal family, including a for each genus showing the estimated native freshwater discussion of each genus, its native distribution, the num- range ( unless indicated other wise) of the genus in North ber of species in each genus, and evidence of polytypy or Amer i ca. For genera with expansive marine ranges, the phylogeographic structure. Contributing authors also in- freshwater and near- shore marine range is given, not the en- cluded a non- native distribution subsection, if informa- tire marine range. Although we took care to make the maps tion was available, outlining introductions outside the as accurate as pos si ble, the scale of the maps and shading of native range and, if known, the e«ects of the taxa as the range obviated pin- point accuracy. Also, for many fishes non- natives. that have been widely introduced, the native range can only Phyloge ne tic relationships. Contributing authors cover all be estimated from often limited historical data. phyloge ne tic hypotheses ( those based on cladistic method- ologies) for the focal family (inter- and intrafamilial) identi- TAXONOMIC CHAPTER ORGANIZATIO N fying, if possi ble, the sister-group of the family and then detailing the relationships of all genera within the family. Each taxonomic chapter contains 13 major sections and Fossil rec ord. Contributing authors summarize the fossil vari ous numbers of subsections. For some families, little history, if any, for the focal family. Minimally the section to nothing may be known about certain sectional and sub- synthesizes information on each known fossil genus in sectional topics, and in those cases, the paucity or lack of the family and the number of extinct fossil species in each information is generally acknowledged. genus. Ages or approximate ages are given when known. Even in a work this large and broad ranging, we came to Morphology. Contributing authors synthesize informa- realize early on that some impor tant topics could not be tion on morphological structures with an emphasis on covered adequately when our focus was largely at the level diversity in morphology across family members and spe- of genus. Hence, contributors did not cover the zoogeogra- cialized, unusual, and unique features. In the introduc- phy of species within each family. The zoogeography of tory subsection, contributing authors describe the general North American freshwater fishes would require another physiognomy of the respective family (e.g., body shape; fin volume to update and reassess information previously syn- shape, type, relative size, and placement; mouth size and thesized on that topic (e.g., Hocutt & Wiley 1986; Mayden placement; scale type, color, and pigmentation patterns).

349-82339_Warren_ch00_1P.indd 16 10/24/19 10:32 PM PREFACE xvii

In other subsections, contributing authors detail unusual The section focuses on topics such as age and size at matu- or specialized external and internal anatomical character- rity; sexual dimorphism; spawning migrations, cues, and istics (e.g., reproductive anatomy, sensory organs, func- sites; pre- spawning and spawning be hav iors; male and tional biology, and eco- morphology). female reproductive allocation; parental care; unusual Ge ne tics. Contributing authors synthesize genetic- based mating systems; and embryo and larval development. studies focusing on topics such as karyology, phylogeogra- Ecol ogy. Contributing authors focus on habitat, diet phy, infraspecific variation, and hybridization and intro- (particularly diet breadth and specializations), predation, gression. They also pre sent studies employing genes or and parasitism but also when pos si ble range across topics gene products to determine phylogeny in the phylogene tic from autecol ogy to the functional importance of individ- relationships section. ual taxa in communities and ecosystems. Physiology. Contributing authors highlight the incredi- Conservation. Contributing authors discuss imperiled ble diversity of physiological traits exemplified by fishes in fishes in the focal family and the likely reasons for their each family. When information is available, the sections decline. Contributing authors incorporate the best avail- include syntheses on tolerances to and e«ects of water able information and summarize the reasons for declines temperature, dissolved oxygen, pH, salinity, and turbidity. or anticipated declines. Contributing authors also highlight other specialized Commercial importance. Contributing authors cover eco- physiological adaptations of each family (e.g., swimming nomic importance and values of taxa in the focal family. per for mance, sensory physiology, chemical ecol ogy, bio- This includes the importance in historic or pre sent com- energetics, and metabolism). mercial fisheries, cultural significance, aquaculture, sport Be hav ior. Contributing authors cover nonreproductive- fisheries, and the aquar ium trade. associated be hav iors in this section. These include behav- Lit er a ture guide. In the final section, contributing au- ioral areas such as aggression, dominance, learning, thors point the reader to major sources of information on memory, migratory and nonmigratory movement, diel activ- the family. In par tic u lar, detailed family, species, or topic- ity, schooling be hav ior, foraging be hav ior, alarm signaling, specific treatments are referenced. patch choice, and any other unique or specialized be hav ior. Reproduction. Contributing authors synthesize features Melvin L. Warren, Jr. of the reproductive cycle, including reproductive be hav iors. Brooks M. Burr

349-82339_Warren_ch00_1P.indd 17 10/24/19 10:32 PM This page intentionally left blank Acknowl edgments

We acknowledge Richard L. Mayden for originating the Katherine Smith, formerly Aquatic Ecologist, O°ce of the core concept of this work and getting it underway. We Chief, Research and Development, USDA Forest Ser vice, thank Vincent Burke, formerly with Johns Hopkins Univer- for substantial and continued support of the project. sity Press, for wise counsel, steady guidance, and encour- agement on diverse matters that arose in finalizing manu- COLOR PLATES AND PHOTO GRAPHS scripts and coordinating with the contributors. Ti«any Gasbarrini and Esther Rodriguez with the Press always We used color drawings by Joe Tomelleri, the premier fish promptly responded to questions concerning myriad details artist in North Amer i ca, to illustrate as many North Ameri- associated with readying manuscripts for publication and can genera as pos si ble. The colors are as seen on the fishes guided the editors through the publication pro cess. Many when they are first removed from the water, and many thanks to Rob Hopkins for designing and painstakingly were drawn when at their peak breeding colors. Joe’s fish preparing the distribution maps and patiently revising portraits are done in Berol Prismacolor pencils, and using them to the satisfaction of the editors and contributors. Our those pencils Joe is renowned for precisely portraying life copyeditor, Liz Radojkovic, was simply superb in attention color, scale and fin- ray counts, and a full spread of the fins. to myriad details of style, consistency, and clarity. We express our appreciation to Joe for granting us a gener- Numerous other individuals worked diligently with ous licensing arrangement for use of his drawings. us to prepare this volume. We appreciate Gayle Hender- For contributing or providing liberal licensing agree- son and Amy Commens- Carson for creating and redraw- ments of copyrighted photo graphs to Volume 2, we are ing numerous figures. Gordon McWhirter, Kenneth Ster- grateful to Ginny Adams; Glynn Aland; Alaska State ling, and Vicki Reithel patiently formatted, proofed, and Library Butler / Dale Photo Collection; American Livebearer cross- referenced numerous drafts of a large and ever- Association; American Society of Ichthyologists and expanding lit er a ture cited as well as in- text references to Herpetologists; Juan Miguel Artigas Azas; Tom Baugh; tables and figures. Mickey Bland assiduously ran down Claude D. Barbour; Michael G. Bennett; Paul Bentzen; often old or obscure lit er a ture and kept the papers coming. Biodiversity Heritage Library; Richard Betancur- R.; Zanethia Choice and Daniel Warren proofed vari ous as- Mickey Bland and Bryan Simmons; John Brill; Nancy pects of text, tables, and figures. Wendell Haag provided Brown- Peterson; Bulletin of Marine Science; Brooks M. guidance on mussel fish- host lit er a ture. Nancy Smith and Burr; California Department of Water Resources; Evan Sonjie Schwartz kindly and adeptly coordinated financial Carson; Robert C. Cashner; Ben Chapman; David Cline; contributions, and Cathy Jenkins and Brenda Marshall Kristin Co«ey; Adam Cohen; Michael Collyer; Steve provided critical logistical support. We also acknowledge Creek; Matthew P. Davis; William P. Davis; Department Ted Leininger, Proj ect Leader, and Susan Adams, Team of Anthropology, Smithsonian Institution; Mauricio De la Leader, Center for Bottomland Hardwoods Research, Maza Benignos; Anthony A. and Alice F. Echelle; Marco Southern Research Station, USDA Forest Ser vice, and Endruweit (aquariophil. org); Eric Engbretson, Jennifer

349-82339_Warren_ch00_1P.indd 19 10/24/19 10:32 PM xx ACKNOWLEDGME NTS

Idol, Sean Landsman, Isaac Szabo, and Paul Vecsei (Eng- Burkhead, Stephen T. Ross, and William N. Roston for bretson Underwater Photography); Danté Fenolio (www granting us and the contributing authors blanket permis- . anotheca . com); Michael Fine; William Fink; Dean Fletcher; sion to use any of their ﬁsh photo graphs. G. Funkner; Dave Gordon; Lance Grande; Andreas Hartl; Fi nally, I (MLW) dedicate this volume to my wife and Gene Helfman; Dean Hendrickson; Michael J. Ghedotti; life partner, Ann Phillippi. She listened to my editorial Adán Gómez- González; Lance Grande; Terry Grande; woes and gently urged me through dark days and was also Charley Grimes; H&H Lure Com pany; A. Hartl; Chad there to share in my triumphs during the editing pro cess, Hood; L. Horth; Samuel Ja«e; W. R. Je«ery; G. I. Jofre; all the while getting our two sons through their educa- Ernest Keeley; Thomas Kline (Alaska Stock . com); Jean tional careers, running our house hold, and carry ing on Krejca; Bernie Kuhajda; Sean Landsman Photography; with her own career in biology and nature outreach to Eric Lang; B. Langerhans; Sean Lema; Robert Liu; Paul V. young minds. My two wonderful sons, Daniel and Mac, Loiselle; Andrés López; John Lundberg; John D. Lyons; Hen- likewise were always there to encourage and support me. rik Malchau; Richard L. Mayden; Caleb McMahan; Carole For our close and internally supportive family, I am most McIvor; Lance Merry; Maurice Mettee; Paul Miano; grateful. Bob Michelson, Photography by Michelson, Inc.; M. Meyer; Missouri Department of Conservation; National FINANCIAL AND IN- KIND SUPPORTERS Park Ser vice; Dave Neely; Douglas Nelson; Heok Hee Ng; Barbara Nicca; Leo Nico; NOAA Photo Library; Claudia P. Alaska Fisheries Science Center, National Marine Fisher- Ornelas- García; Ohio Department of Natu ral Resources; ies Ser vice, NOAA Desert Fishes Council Michael Patrick O’Neill / Alamy; Tom Palmer; Seth Patter- Center for Bottomland Hardwoods Research, Southern son; Kyle R. Piller; Pat Rakes; René Reyes; A. Riggs; Fritz Research Station, USDA Forest Ser vice Rohde; Rocio Rodiles; Paddy Ryan; Yoshitaka Sakakura; Museum of Southwestern Biology, University of New Manuel M. Salazar; Joe Scanlan; J. R. Shute; Bernard Siet- Mexico man; Sjh123 at En glish Wikipedia; Scott Simmons; Smith- North American Native Fishes Association sonian Institution; Garold Sneegas; F. F. Snelson, Jr.; Springer Oklahoma State University Science+Business Media; Je« Stewart; Ulrike Strecker; Roanoke College Colette St Mary; Roger A. Tabor; A. Tatarenkov; D. Scott Robert C. Cashner Taylor; T. Taylor and J. Jackson; Nate Tessler; The Wilder- Southeastern Fishes Council ness Medical Society; Cheryl Theile; Matt Thomas; Uland Southern Division of the American Fisheries Society Thomas; Marc Thorpe / Marc Thorpe Guiding Ser vice; Tennessee Aquar ium Conservation Institute Bob Trapani; University of Michigan Museum of Zoology; University of Iowa Stephano Valdesalici; Frans Vermeulen; Stephen J. Walsh; University of Toronto Mel Warren; Shawn A. Webb; E. Wieand; Scott Wickard; Southeast Ecological Science Center, United States Geo- Wilderness Medical Society; Joe Worthem courtesy of In- logical Survey vitation Oxford magazine; and Patricia Wright. Likewise, Research and Development, O°ce of the Chief, USDA we gratefully acknowledge Anthony Terceira, Noel M. Forest Ser vice

349-82339_Warren_ch00_1P.indd 20 10/24/19 10:32 PM