Ecology of North American Freshwater Fishes. Stephen T. Ross Ph. D.
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for this work, and many others, through his monumental book Patterns in Freshwater Fish Ecology. I especially appreciate Chuck Crumly at the University of California Press for inviting me to write a textbook on the ecology of North American freshwater fishes. I thank Lynn Meinhardt, also of the University of California Press, for her helpful comments and encouragement throughout.
During my work on this book I have benefited greatly from the support of my colleagues at the University of New Mexico, Museum of Southwestern Biology and the Department of Biology, particularly Tom Turner, Manuel Molles, and Alexandra (Lex) Snyder. I thank Tom for his helpful comments and suggestions during the course of this project, and for the opportunity to present some of the chapter contents for discussions with the Turner lab group. Manuel, a textbook author himself, has been more than patient in guiding me through the intricacies of textbook writing and in vetting my use of ecological terms and concepts. I appreciate Lex, who has always been supportive and has made resources of the Division of Fishes readily available to me. I am grateful to Steve Platania, Rob Dudley, Howard Brandenburg, and Mike Farrington of the Museum of Southwestern Biology (MSB) and American Southwest Ichthyological Researchers (ASIR) for providing help with literature and for giving me access to unpublished data. Howard Brandenburg also provided slides of the San Juan River, and I thank both Howard and David Propst, formerly of New Mexico Game and Fish, for giving me permission to use Howard’s outstanding drawings of New Mexico fishes. I also thank Tom Kennedy for generously allowing me to use his photographs of southwestern fishes and Trevor Krabbenhoft for providing larval fish pictures. The faculty and staff of the Interlibrary Loan Department of the University of New Mexico have provided exceptional help throughout the duration of this book. Indeed, without their assistance this book would not have been possible.
I am grateful to colleagues around the country who have given such tremendous support, sharing information on their research and findings. I thank John Baker always for his enthusiastic help, particularly on stickleback biology, and Ron Bliesner, of Keller-Bliesner Engineering, for allowing me to use his GIS analysis of the San Juan River. Other colleagues have also been generous in allowing me to use their photographs of fishes and habitats, although page limitations precluded using many of the images. For this I thank John Baker and Anna Mazzarella (Clark University), Peter Bisson (Pacific Northwest Research Station, USDA Forest Service), Mollie Cashner (Southeastern Louisiana University), Katie May Laumann (Virginia Institute of Marine Science), Gary Meffe (University of Florida), James Morel (Navajo Nation Department of Fish and Wildlife), and W. Todd Slack and Steven G. George (Waterways Experiment Station, USACE).
I thank Tim Modde (USFWS) for including me on research expeditions on the Yampa and Green rivers of the Colorado system years ago, and for providing input on the Razorback Sucker and Colorado Pikeminnow. I am also grateful to Marlis and Mike Douglas for including me on a research expedition through the Grand Canyon of the Colorado River.
I am forever indebted to colleagues who devoted the time and effort to review entire chapters: Chapter 7, Trevor Krabbenhoft and Evan Carson, University of New Mexico; Chapter 8, Trevor Krabbenhoft and Tracy Diver, University of New Mexico; Chapter 9, Eric Charnov, University of New Mexico, and David Heins, Tulane University; Chapter 10, Trevor Krabbenhoft; Chapter 11, Tom Turner, University of New Mexico; Chapter 12, Chet Rakocinski, University of Southern Mississippi; Chapter 13, Mike Farrington, University of New Mexico and ASIR; Chapter 14, Megan Osborne, University of New Mexico; Chapter 15, Corey Krabbenhoft and Nathan Franssen, University of New Mexico.
As in all my previous endeavors throughout my career in biology, work on this book has been greatly aided, indeed made possible, by Yvonne Ross, my wife, editor, and best friend for more than 40 years, who read and commented on the entire text. Our son, Derek G. Ross (Auburn University), has helped tremendously by leading me into twenty-first-century technology in my writing, bibliographic research, and in organizing information. He also lent his expertise in technical communication, scientific rhetoric, and ethics by contributing one of the text boxes. Finally, travels across North America for habitat pictures as well as day-to-day writing have all been helped by the constant supervision and diversion of Bergen, our indefatigable canine companion.
PART ONE
Faunal Origins, Evolution, and Diversity
Fishes represent the most diverse and species-rich group of craniate organisms. In this part, the focus is on introducing the freshwater fish faunas of the world and on the unique nature of the North American freshwater fish fauna (Chapter 1). Chapters 2 and 3 deal with the very basic issues of where the North American freshwater fish fauna came from, how long various fish lineages have occurred in North America, and how the fauna has been shaped by large scale geologic and climatic events.
A useful way of thinking about why certain fish species occur in particular regions, water bodies, and local habitats is to envision an ancestral fauna being shaped by a series of filters. Some filters can be stronger than others and, of course, some taxa might be better at surmounting the challenges imposed by the filters. The broadscale filters are the concern of Part 1 (see the following figure), the end result being an understanding of the origins and ages of North American regional fish faunas. Subsequent parts will move through smaller scale, more local filters, leading to an understanding of factors shaping the occurrence of fishes in local habitats such as a particular pond, lakeshore, riffle, or run.
A conceptual model of the formation of fish assemblages through progressive loss (i.e., filtration) and through the progressive addition (i.e., speciation) of lineages. Adapted in part from Smith and Powell (1971); Poff (1997); Matthews (1998); Tonn et al. (1990); Rahel (2002); Ross and Matthews (in press).
ONE
Introduction
CONTENTS
Distribution of Fresh Water: Global Patterns
North American Freshwater Fishes
Patterns of North American Diversity
UNDERSTANDING FISH ECOLOGY IS both exciting and challenging. Fishes have the greatest diversity among the craniate organisms, composing more than half of all extant species, with an estimated 27,977 species of fishes worldwide (Nelson 2006), and with the descriptions of “new” (newly documented) species continuing at approximately 200 species per year (Eschmeyer 1998). As further emphasis of fish diversity, unlike other recognized groups within the phylum Chordata (amphibians, reptiles, birds, and mammals), organisms commonly referred to as fishes comprise five living classes (hagfishes, Myxini; lampreys, Petromyzontidae; sharks and rays, Chondrichthyes; ray-finned fishes, Actinopterygii; and lobe-finned fishes, Sarcopterygii). Three of these groups, petromyzontids, chondrichthyans, and actinopterygians, are represented in North American fresh water. The Myxini are restricted to the marine environment, whereas the fishes included within the class Sarcopterygii occur in the marine environment (Coelacanthimorpha, coelacanths) or fresh waters of Australia, Africa, and South America (Dipnoi, lungfishes). Tetrapods, including, of course, readers of this book, are also included in the Sarcopterygii (Nelson 2006).
Despite the abundance of organisms to study, the aquatic environment provides challenges to work in, making ecological studies difficult or even dangerous. Especially in many North American streams, visual observations are often not possible because of high flows or turbidity, necessitating indirect approaches, such as seining, electrofishing, or using electronic tags, to infer information on fish habitat use, behavior, and interactions.
This is nonetheless an exciting, as well as challenging, time to write about North American freshwater fish ecology. Modern computers and the rapidly increasing quality of electronic libraries and data searches have made the literature much more available, and the rate of publications has greatly accelerated. In