Volume 20 Number 2 October 2017
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FRESHWATER MOLLUSK BIOLOGY AND CONSERVATION THE JOURNAL OF THE FRESHWATER MOLLUSK CONSERVATION SOCIETY VOLUME 20 NUMBER 2 OCTOBER 2017 Pages 33-58 oregonensis/kennerlyi clade, Gonidea angulata, and A Revised List of the Freshwater Mussels (Mollusca: Margaritifera falcata Bivalvia: Unionida) of the United States and Canada Emilie Blevins, Sarina Jepsen, Jayne Brim Box, James D. Williams, Arthur E. Bogan, Robert S. Butler, Donna Nez, Jeanette Howard, Alexa Maine, and Kevin S. Cummings, Jeffrey T. Garner, John L. Harris, Christine O’Brien Nathan A. Johnson, and G. Thomas Watters Pages 89-102 Pages 59-64 Survival of Translocated Clubshell and Northern Mussel Species Richness Estimation and Rarefaction in Riffleshell in Illinois Choctawhatchee River Watershed Streams Kirk W. Stodola, Alison P. Stodola, and Jeremy S. Jonathan M. Miller, J. Murray Hyde, Bijay B. Niraula, Tiemann and Paul M. Stewart Pages 103-113 Pages 65-70 What are Freshwater Mussels Worth? Verification of Two Cyprinid Host Fishes for the Texas David L. Strayer Pigtoe, Fusconaia askewi Erin P. Bertram, John S. Placyk, Jr., Marsha G. Pages 114-122 Williams, and Lance R. Williams Evaluation of Costs Associated with Externally Affixing PIT Tags to Freshwater Mussels using Three Commonly Pages 71-88 Employed Adhesives Extinction Risk of Western North American Freshwater Matthew J. Ashton, Jeremy S. Tiemann, and Dan Hua Mussels: Anodonta nuttalliana, the Anodonta Freshwater Mollusk Biology and Conservation ©2017 ISSN 2472-2944 Editorial Board CO-EDITORS Gregory Cope, North Carolina State University Wendell Haag, U.S. Department of Agriculture Forest Service Tom Watters, The Ohio State University EDITORIAL REVIEW BOARD Conservation Jess Jones, U.S. Fish & Wildlife Service / Virginia Tech University Ecology Ryan Evans, Kentucky Department of Environmental Protection, Division of Water Michael Gangloff, Appalachian State University Caryn Vaughn, Oklahoma Biological Survey, University of Oklahoma Freshwater Gastropods Paul Johnson, Alabama Aquatic Biodiversity Center Jeff Powell, U.S. Fish & Wildlife Service, Daphne, Alabama Jeremy Tiemann, Illinois Natural History Survey Reproductive Biology Jeff Garner, Alabama Division of Wildlife and Freshwater Fisheries Mark Hove, Macalester College / University of Minnesota Survey/Methods Heidi Dunn, Ecological Specialists, Inc. Patricia Morrison, U.S. Fish & Wildlife Service Ohio River Islands Refuge David Strayer, Cary Institute of Ecosystem Studies Greg Zimmerman, Enviroscience, Inc. Systematics/Phylogenetics Arthur Bogan, North Carolina State Museum of Natural Sciences Daniel Graf, University of Wisconsin-Stevens Point Randy Hoeh, Kent State University Toxicology Thomas Augspurger, U.S. Fish & Wildlife Service, Raleigh, North Carolina Robert Bringolf, University of Georgia John Van Hassel, American Electric Power Teresa Newton, USGS, Upper Midwest Environmental Sciences Center Freshwater Mollusk Biology and Conservation 20:33–58, 2017 Ó Freshwater Mollusk Conservation Society 2017 REGULAR ARTICLE A REVISED LIST OF THE FRESHWATER MUSSELS (MOLLUSCA: BIVALVIA: UNIONIDA) OF THE UNITED STATES AND CANADA James D. Williams1*, Arthur E. Bogan2, Robert S. Butler3,4,KevinS.Cummings5, Jeffrey T. Garner6,JohnL.Harris7,NathanA.Johnson8, and G. Thomas Watters9 1 Florida Museum of Natural History, Museum Road and Newell Drive, Gainesville, FL 32611 USA 2 North Carolina Museum of Natural Sciences, MSC 1626, Raleigh, NC 27699 USA 3 U.S. Fish and Wildlife Service, 212 Mills Gap Road, Asheville, NC 28803 USA 4 Retired. 5 Illinois Natural History Survey, 607 East Peabody Drive, Champaign, IL 61820 USA 6 Alabama Division of Wildlife and Freshwater Fisheries, 350 County Road 275, Florence, AL 35633 USA 7 Department of Biological Sciences, Arkansas State University, State University, AR 71753 USA 8 U.S. Geological Survey, Wetland and Aquatic Research Center, 7920 NW 71st Street, Gainesville, FL 32653 USA 9 Museum of Biological Diversity, The Ohio State University, 1315 Kinnear Road, Columbus, OH 43212 USA ABSTRACT We present a revised list of freshwater mussels (order Unionida, families Margaritiferidae and Unionidae) of the United States and Canada, incorporating changes in nomenclature and systematic taxonomy since publication of the most recent checklist in 1998. We recognize a total of 298 species in 55 genera in the families Margaritiferidae (one genus, five species) and Unionidae (54 genera, 293 species). We propose one change in the Margaritiferidae: the placement of the formerly monotypic genus Cumberlandia in the synonymy of Margaritifera. In the Unionidae, we recognize three new genera, elevate four genera from synonymy, and place three previously recognized genera in synonymy. We recognize for the first time two species (one native and one nonindigenous) in the Asian genus Sinanodonta as occurring in North America. We recognize four new species and one subspecies and elevate 21 species from synonymy. We elevate 10 subspecies to species status and no longer recognize four subspecies. We change common names for five taxa, correct spelling for eight species, and correct the date of publication of original descriptions for four species. KEY WORDS: Unionidae, Margaritiferidae, taxonomy, systematics, nomenclature, mussel scientific names, mussel common names INTRODUCTION and Canada. Much of this interest was brought about by passage of the U.S. Endangered Species Acts of 1966, 1969, During the past 50 yr, there has been considerable interest and 1973 and the Canadian Species at Risk Act of 2002. These in freshwater mussels (order Unionida) in the United States legislative actions and the environmental movement that accompanied them focused conservation attention on all *Corresponding Author: [email protected] animals and plants, as well as their habitats. This in turn led 33 34 WILLIAMS ET AL. to assessment of species conservation status and the iczn.org/iczn/index.jsp). Species mentioned in the text, but not development of faunal lists for many states and provinces. included in Table 2, have author and date of publication The task of developing species lists was difficult for most following the name. Author and date of publication for all invertebrates, including mussels, because so little attention had other species are given in Table 2. been given to the study of their biology, ecology, and Mussel common names follow Turgeon et al. (1998) with systematics. In 1970, only six U.S. states had recent lists or minor exceptions, but they are capitalized as is now the books covering their mussel fauna. The first modern attempt to practice for many other animal groups (e.g., birds, reptiles, provide a comprehensive list of freshwater mussels of North amphibians, fishes). Capitalization of common names helps America was published by Burch (1973, 1975). avoid confusion by identifying standardized common names. The first comprehensive list of freshwater mussels of the For example, reference to a ‘‘fragile papershell’’ could apply to United States and Canada was compiled in Turgeon et al. several thin-shelled species, but the capitalized ‘‘Fragile (1988) and revised a decade later (Turgeon et al. 1998). Papershell’’ is unambiguously recognized as the common Williams et al. (1993) was another important resource during name for Leptodea fragilis. We note and explain other this period; although mainly an assessment of species instances where we changed common names from those of conservation status, this paper also provided a comprehensive Turgeon et al. (1998) or where recognition of previously and widely used species list similar to those of Turgeon et al. unrecognized species necessitated creation of a new common (1988, 1998). These lists standardized and provided taxonomic name. stability to mussel common and scientific names to an extent We provide a rationale for and discussion of all taxonomic that was previously unavailable. However, systematic taxon- changes in the following accounts for each family and genus omy of mussels was poorly known at that time, and and in Table 2. There is a degree of uncertainty and classifications at all taxonomic levels were based largely on subjectivity in our revised list that is unavoidable given our concepts from the early 1900s. still imperfect understanding of mussel systematics. We Since publication of Turgeon et al. (1988, 1998) and attempted to reconcile divergent views regarding mussel Williams et al. (1993), many studies have refined our systematics based on our assessment of the strength of understanding of mussel systematic taxonomy. Several major evidence for these views. In cases where evidence did not publications have addressed systematic relationships within allow reconciliation, we attempted to provide a plausible the class Bivalvia, including the order Unionida (Bieler et al. conclusion based on our professional judgment and experi- 2010; Carter et al. 2011; Bolotov et al. 2016; Araujo et al. ence; these conclusions were based on consensus among the 2017; Combosch et al. 2017). Major studies specific to the authors to the extent possible. Unionida include Graf and O´ Foighil (2000), Hoeh et al. Subspecies is a taxonomic category applied to populations (2001, 2002, 2009), Roe and Hoeh (2003), Campbell et al. that are morphologically distinct and geographically separated (2005), Walker et al. (2006), Graf and Cummings (2007, but that exhibit intergradation in contact zones (Mayr et al. 2017), Cummings and Graf (2010), and Campbell and 1953; Gilbert 1961). We evaluated morphological and Lydeard (2012a, 2012b).