Determination of the Systematic Position And

Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 1

Determination of the systematic position and relationships of the Atlantic Pigtoe, Fusconaia masoni (Conrad, 1834) (Mollusca: Bivalvia: Unionidae) with distributions in Virginia, North and South Carolina, and Georgia.

Final Report

Arthur E. Bogan1, Morgan Raley2 and Jay Levine2

1North Carolina State Museum of Natural Sciences, Research Laboratory, 4301 Reedy Creek Road, Raleigh, NC 27607 Tel. 919-733-7450 ext 753 [email protected] 2North Carolina State University, College of Veterinary Medicine, Raleigh, NC

USFWS Contract

1 June 2003

Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 2

ABSTRACT North America is home to the most diverse freshwater bivalve fauna in the world. However, it also has the distinction of having lost the most species to extinction, 32 species, than any other area in the world. Today, 75% of the native North American freshwater bivalve fauna (Unionoida) is imperiled. The rivers of the southern Atlantic Slope are poorly studied and the molluscan fauna is no exception. If coastal rivers are actually biogeographic islands, then many of the described taxa found in these isolated coastal rivers may represent endemic species. We have examined the sequence of a section of the first subunit of cytochome oxidase c gene (COI), a mitochondrial gene, from a series specimens of the Atlantic Pigtoe to determine if this is a single species and if the species has been placed in the correct genus. The conclusion is that Fusconaia masoni is a good species and belongs in the genus Fusconaia. INTRODUCTION Johnson (1970) has provided the most recent review of freshwater mussels occurring along the southern Atlantic Slope Region of eastern North America. The Atlantic Pigtoe, Fusconaia masoni (Conrad, 1834) was described from the Savannah River, Augusta, Richmond County, Georgia (Conrad, 1834:34, pl. 5, fig. 2). The Atlantic Pigtoe ranges from the Ogeechee River Basin in Georgia, north to the James River Basin in Virginia (Johnson, 1970). It should be noted that Johnson (1970) synonymized Lexingtonia subplana under Fusconaia masoni in his treatment of the species. Fusconaia masoni is known historically from all of the coastal river basins in North Carolina except the Cooper-Santee and the Waccamaw River basins (Fuller, 1977; Bogan 2002) and extends from the Chowan River basin of southern Virginia in the north (Fuller, 1975), south to the Ogeechee River (Fuller, 1973, Burch 1975). The early work on biochemical analyses of Unionidae Davis and Fuller (1981), Davis, et al. (1981), Davis (1983, 1984), Kat (1983), Kat and Davis (1984), and Davis and Mulvey (1993) attempted to address the relationships among some of the Atlantic Slope taxa but did not deal with either Fusconaia masoni or Lexingtonia subplana. Fuller (1971) was the first to place Unio masoni in the genus Fusconaia and Fuller (1973) fully documented the evidence for this generic assignment. Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 3

This species is not known from the limited Pleistocene Atlantic Slope unioniod faunal record. The reported faunas are located too far north to include this species (Bogan et al. 1989; Bogan and Grady, 1991). The purpose of this project is to determine the taxonomic status of F. masoni and its generic placement using gene sequence data. SYNONYMY Fusconaia masoni (Conrad, 1834); Atlantic Pigtoe Unio masoni Conrad, 1834; Conrad, 1834:34, pl. 5, fig. 2 [Savannah River, Augusta, Richmond County, GA]; Simpson, 1900:709; Simpson, 1914:613-614 Margarita (Unio) masoni (Conrad, 1834); Lea, 1836:33 Margaron (Unio) masoni (Conrad, 1834); Lea, 1852:34 Elliptio (Elliptio) masoni (Conrad, 1834); Frierson, 1927:25; Haas, 1969:216-217 Pleurobema (Lexingtonia) masoni (Conrad, 1834); Johnson, 1970:301-303, pl. 2: figs. 3-10; Burch 1973:16, fig. 35 Fusconaia masoni (Conrad, 1834); Fuller, 1971:142, 143; 1972:72-72; 1973:105-117; 1977:177-178

Unio subplanus Conrad, 1837; Conrad, 1837:73, pl.41, fig.1 [North = Maury River, a branch of the James River, Lexington, Rockbridge Co. Virginia]; Simpson, 1900:720, notes distribution North Carolina and Virginia. Pleurobema subplanum (Conrad, 1837); Simpson 1914:804-805 Lexingtonia subplana (Conrad, 1837); Ortmann, 1914:29-30; Haas, 1969:249 Pleurobema (Lexingtonia) subplanum (Conrad, 1834): Frierson, 1927:44 Pleurobema (Lexingtonia) masoni (Conrad, 1834); Johnson, 1970:301-303 [in part]; Burch, 1973:16

Unio pumilus Lea, 1838; Lea, 1838:23, pl. 7, fig. 17 [Black River, tributary to the Neuse River on the road to Fayetteville from Smithfield, Johnston County, NC; Fuller 1972:72-73; corrected the type locality to: “Black River [Cape Fear River system: South River drainage], on the road to Fayetteville from Smithfield [=about 5 mi. NE Erwin and about 4 mi. ESE Coats, Harnett Co.], North Carolina”] Margarita (Unio) pumilus (Lea, 1836); Lea, 1836:20 [nomen nudum] Margarita (Unio) pumilus (Lea, 1838); Lea, 1838:17 Margaron (Unio) pumilus (Lea, 1838); Lea, 1852:24 Quadrula (Fusconaia) pumila (Lea, 1838); Simpson 1900:785; Simpson 1914:868-869 Pleurobema (Lexingtonia) subplanum (Conrad, 1834); Frierson, 1927:44 [in part] Lexingtonia subplana (Conrad, 1834); Haas, 1969:249 [in part] Pleurobmea (Lexingtonia) masoni (Conrad, 1834); Johnson, 1970:301-303 [in part]

Unio utriculus Lea, 1845; Lea, 1845:164 [North Carolina] Tenn River Basin? Johnson 1970:300 Margaron (Unio) utriculus (Lea, 1845); Lea, 1870:46 Quadrula (Fusconia) utriculus (Lea, 1845); Simpson, 1900:785 lists as North Carolina but notes the type from Arkansas and not from North Carolina!!!! Pleurobema utriculum (Lea, 1845); Simpson, 1914:787 still lists type locality as North Carolina. Pleurobema (Lexingtonia) utriculum (Lea, 1845); Frierson, 1927:45 Lexingtonia utricularis (Lea, 1845); Haas, 1969:249 [North Carolina]

Unio merus Lea, 1852; Lea, 1852:260, pl. 15, fig. 10 [Abbeville District, Savannah River drainage, SC; Simpson, 1900:708; Simpson, 1914:612-613 Margaron (Unio) merus (Lea, 1852); Lea, 1870:54 Elliptio (Elliptio) masoni (Conrad, 1834); Frierson, 1927:25; Haas, 1969:216 [in part] Pleurobmea (Lexingtonia) masoni (Conrad, 1834); Johnson, 1970:301-303 [in part]

Unio striatulus Lea, 1857; Lea, 1857:86; Lea, 1862:55, pl. 2, fig. 202 [Roanoke River, Weldon, Halifax County, NC]; Simpson, 1900:719 Pleurobema striatulum (Lea, 1857); Simpson, 1914:788-789 Pleurobema (Lexingtonia) subplanum (Conrad, 1834); Frierson, 1927:44 Lexingtonia subplana (Conrad, 1834); Haas, 1969:249 [in part] Pleurobmea (Lexingtonia) masoni (Conrad, 1834); Johnson, 1970:301-303 [in part] Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 4

Unio castus Lea, 1860; Lea, 1860:306; Lea: 1860:349; pl. 57, fig. 202 [South Carolina] Margaron (Unio) castus (Lea, 1860); Lea, 1870:40 Unio merus Lea, 1852; Simpson, 1900:708 [in part]; Simpson 1914:612-613 [in part] Pleurobema (Lexingtonia) castus (Lea, 1860); Frierson, 1927:45 Lexingtonia casta (Lea, 1860); Haas, 1969:249 Pleurobmea (Lexingtonia) masoni (Conrad, 1834); Johnson, 1970:301-303 [in part]

Unio brimleyi S.H. Wright, 1897; Wright, 1897:138 [Walnut Creek of Neuse River, Raleigh, Wake County, NC]; Simpson, 1900: 720 Pleurobema brimleyi (S.H. Wright, 1897): Simpson 1914:805 Pleurobema (Lexingtonia) subplanum (S.H. Wright, 1897); Frierson, 1927:44 [in part] Lexingtonia subplana (Conrad, 1834); Haas, 1969:249 [in part] Pleurobema (Lexingtonia) masoni (Conrad, 1834); Johnson, 1970:301-303 [in part]

Unio castus Simpson, 1900 [non Lea, 1860]; Simpson, 1900:708 Elliptio (Elliptio) masoni (Conrad, 1834); Frierson, 1927:25; Haas, 1969:217 [in part]

Unio merus Simpson, 1900 [non Lea]; Simpson, 1900: Pleurobema (Lexingtonia) castum Lea, 1860; Frierson, 1927:45 Comments: Unio utriculus is included here as it continues to be listed as from North Carolina and has been historically used as a member of the South Atlantic Slope fauna, but in fact is probably a synonym of an Interior Basin species. Fuller (1971:72) noted “It will be impossible to construct the correct synonymies of Fusconaia masoni and Lexingtonia subplana until their ranges are fully understood.” Johnson (1970) used the genus Pleurobema and included Lexingtonia Ortmann, 1914 as a subgenus and included P. (L.) collina (Conrad, 1836) and P. (L.) masoni. He included the type species of Lexingtonia, Unio subplanus Conrad, 1837 as a junior synonym of P. (L.) masoni. Lexingtonia and L. subplana have been recognized as a valid separate genus and species by Ortmann (1914), Burch (1975), and Turgeon et al. (1988, 1998). Simpson (1900:720) noted the type locality for Unio subplanus was Virginia but list the distribution as occurring in North Carolina and Virginia. It is clear that many people in the past have followed Ortmann (1914) in recognizing the animal in the James River basin as distinct from that which ranged from the Chowan River south to the Ogeechee River basin. The question of the position and relationships of subplana can only be answered with the inclusion of specimens of Lexingtonia subplana in the DNA analyses with Fusconaia masoni. Lexingtonia subplana is included in the synonymy only for completeness and to show the historic confusion of the relationships of the taxa described from the South Atlantic Coast region. METHODS Specimens and Vouchers Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 5

Whenever possible, two specimens representing each name were sequenced. Outgroup species were chosen based on the results of Lydeard et al. (1996; 2000). In particular, representative species from Pleurobema, Fusconaia, and Lexingtonia were included to test the placement of F. masoni. Voucher specimens are deposited at the North Carolina State Museum of Natural Sciences (NCSM) (see Table 1). DNA Extraction, Amplification, and Sequencing Specimens and sequences were obtained from GenBank or collected from locales listed in Table 1. Foot or mantle tissue was isolated and DNA extracted using the Qiamp Tissue Kit (Qiagen) following the manufacturer's recommendation. DNA integrity was assessed visually on 1% agarose gels. PCR amplification of the partial cytochome c oxidase subunit I (COI) gene followed the methods of Folmer, et al. (1994). PCR products were purified using the QIAquick PCR Purification Kit (Qiagen) following the manufacturer's recommendation. DNA sequencing was performed using the same primers used for PCR (LC01490 & HC02198) following the recommended protocol with the ABI BigDye v. 3.0 sequencing kit (Applied Biosystems). Sequences were purified for sequencing using the Qiagen DyEx Spin Kit (Qiagen) and visualized using an ABI 377 automated sequencer (Applied Biosystems). Sequences were aligned visually and assembled into a PAUP matrix for phylogenetic analysis. Phylogenetic analysis was attempted with PAUP*4.0 b10 (Swofford, 2002) on a Power Macintosh G3. Heuristic searches were attempted with 100 replicates using tree bisection-reconnection (TBR) branch swapping algorithm. Additional parameters included 1) only trees of minimal length retained, and 2) branches of zero length collapsed. It should be noted, however, that due to the constraints of the computer used in the analysis (memory, processor speed, etc.) that once 17,500 trees were saved, parsimony analysis proceeded without reaching a most parsimonious resolution. We have decided to forgo any further parsimony analysis until a suitable machine or algorithm may be found with which to analyze this dataset. Therefore Neighbor-Joining trees are presented as preliminary results for this analysis. RESULTS Preserved specimens of Lexingtonia subplana from Ohio State University, Museum of Zoology, were examined and an attempt was made to extract DNA from the preserved Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 6 tissue. Jeanne Serb (Pers. Comm.) was unable to extract usable DNA from those specimens. We have just received tissue samples from two specimens form the University of Georgia Museum of Zoology, of F. masoni collected in 1991 from Williamson Swamp Creek, Ogeechee River Basin in Georgia (Alderman, 1991). The specimens are being prepared for extraction at this time. Nothing is known of their preservation history. We have been able to obtain live specimens from the Nottaway River, Chowan River Basin, and Moccasin Creek and Turkey Creek, Neuse River Basin. There are trips planned to try and collect live specimens of F. masoni from Aarons Creek, a tributary to the Dan River, Roanoke River Basin. We have not been able to obtain specimens of L. subplana to date. All of the specimens of F. masoni examined to date and included in Figure 1 form a monophyletic clade allied with the clade containing Fusconaia flava, type species of the genus Fusconaia. This would support the claim that the species U. masoni belongs in the genus Fusconaia. This analysis compared F. masoni with conchologically similar Pleurobema georgiana, Fusconaia barnesiana as well as Lexingtonia dolabelloides from the Duck River, Tennessee, the supposed sister species of L. subplana. Lexingtonia dolabelloides clusters with F. barnesiana but is separated from the rest of the Fusconaia clade by the Pleurobema clade (Figure 1). DISCUSSION Ortmann (1914), Fuller (1971) and Burch (1975) listed the distribution of Lexingtonia subplana as restricted to the James River Basin, Virginia while F. masoni was found from the Chowan River basin south to the Ogeechee River Basin. Fuller (1973) “proposed that precursors of modern Atlantic drainage Fusconaia and Lexingtonia migrated from the ancestral New River drainage during its form confluence with the Roanoke River System.” Our analysis has established Fusconaia masoni is a good species and belongs in the genus Fusconia based on the COI sequence data. This supports the placement based on the species use of all four gills as marsupia and the coloration of the marsupia during the development of the embryos, characteristics of the genus Fusconia (Fuller, 1973). Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 7

Future work on this species needs to include more specimens from other river basins as well as if at all possible specimens of L. subplana from the James River basin. ACKNOWLEDGMENTS We would like to thank the following people for assistance in the field: Margaret Barfield,; Chris Eads, NCSUCVM and field crew, Sean N. Bogan, Ian J.G. Bogan, Cynthia M. Bogan, Cary NC; Tim Savidge NCDOT, John Alderman, NCWRC, John Fridell USFWS Asheville, NC; Judy Johnson-Ratcliff NCWRC; Valeria Rice, NCSM, David Davenport, NCSM; E. Keferl, GA, W. Todd Slack, and several fellow collectors from MMNS LITERATURE CITED Alderman, J.M. 1991. Status survey for the Atlantic Pigtoe (Fusconaia masoni) in Georgia. Nongme and Endangered Wildlife Program. Division of Wildlife Management, North Carolina Wildlife Resources Commission, 6 pp, 49 maps. Bogan, A.E. 2002. Workbook and key to the freshwater bivalves of North Carolina. North Carolina Museum of Natural Sciences, Raleigh, NC. 101 pp, 10 color plates. Bogan, A.E. and F.V. Grady. 1991. Two Pleistocene Molluscan faunas from Eastern West Virginia. In: Beamers, bobwhites and blue-points: tributes to the career of Paul W. Parmalee. pp. 189-213. J.R. Purdue, W.E. Klippel and B.W. Styles (eds.). Illinois State Museum Scientific Papers, Vol 23, Springfield and The University of Tennessee, Department of Anthropology Report of Investigations No. 52, Knoxville. Bogan, A.E., E.E. Spamer, G.C. Manville, W.B. Gallagher and A.J. Cain. 1989. Preliminary reexamination of the Fish House Local Fauna and Flora (Pleistocene), Pennsauken, Camden County, New Jersey. Mosasaur 4:111-126. Burch, J. B. 1973. Freshwater Unionacean clams (Mollusca: Pelecypoda) of North America. Biota of Freshwater Ecosystems. Identification Manual 11. U.S. Environmental Protection Agency. 176 pp. Burch, J. B. 1975a. Freshwater Unionacean clams (Mollusca: Pelecypoda) of North America. Rev. ed. Malacological Publications. Hamburg, Michigan. 204 pp. Conrad, T. A. 1834. New freshwater shells of the United States, with coloured illustrations; and a monograph of the genus Anculotus of Say; also a synopsis of the American naiades. J. Dobson, 108 Chestnut Street, Philadelphia, Pennsylvania. 1–76, 8 pls. Conrad, T. A. 1835b–1840. Monography of the Family Unionidae, or naiades of Lamarck, (fresh water bivalve shells) or North America, illustrated by figures drawn on stone from nature. J. Dobson, 108 Chestnut Street, Philadelphia, Pennsylvania. 1835, 1:1–12, [pp. 13–16 not published], pls. 1– 5; 1836, 2:17–24, pls. 6–10; 1836, 3:25–32, pls. 11–15; 1836, 4:33–40, pls. 16–19; 1836, 5:41–48, pls. 21–25; 1836, 6:49–56, pls. 26–30; 1836, 7:57–64, pls. 32–36; 1837, 8:65–72, pls. 36–40; 1837, 9:73–80, pls. 41–45; 1838, 10:81–94, 2i, pls. 46–51; 1838, 11:95–102, pls. 52–57; 1840, 12:103–110, pls. 58–60; [1840], 13:111–118, pls. 61–65. Davis, G.M. 1983. Relative roles of molecular genetics, anatomy, morphometrics and ecology in assessing relationships among North American Unionidae (Bivalvia). Pp. 193-222. IN: Systematics Association Special Volume No. 24. Protein polymorphism: adaptive and taxonomic significance. G.S. Oxford and D. Rollinson (editors). Academic Press. Davis, G.M. 1984. Genetic relationships among some North American Unionidae: sibling species, convergence, and cladistic relationships. Malacologia 25:629-648. Davis, G.M. and S.L.H. Fuller. 1981. Genetic relationships among Recent Unionacea (Bivalvia) of North America. Malacologia 20(2):217-253. Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 8

Davis, G.M. W.H. Heard, S.L.H. Fuller and C. Hesterman. 1981. Molecular genetics and speciation in Elliptio and it relationships to other taxa of North American Unionidae (Bivalvia). Biological Journal of the Linnean Society 15(2):131-150. Davis, G.M. and M. Mulvey. 1993. Species status of Mill Creek Elliptio. SRO-NERP-22. A publication of the Savannah River Site, 58 pp. Folmer, O., W.R. Hoeh, M.B. Black, R.L. Vrijenhoek. 1994. DNA primers for amplification of mitochondrial cytochrome C oxidase subunit I from metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294-299. Frierson, L.S. 1927. A classified and annotated check list of the North American naiades. Baylor University Press, Waco, Texas. 111 pp. Fuller, S.L.H. 1971. A brief field guide to the fresh-water mussels (Mollusca: Bivalvia: Unionacea) of the Savannah River System. The ASB Bulletin 18(4):137-146. Fuller, S.L.H. 1972. Type locality of Unio pumilus Lea, 1838 (Unionidae). The Nautilus 86(2-4):72-73. Fuller, S.L.H. 1973. Fusconia masoni (Conrad, 1834) (Bivalvia: Unionacea) in the Atlantic Drainage of the Southeastern United States. Malacological Review 6:105-117. Fuller, S.L.H. 1975. Fresh-water mussels (Mollusca: Bivalvia) in the Chowan river System of Virginia. The ASB Bulletin 22(2):54. Fuller, S.L.H. 1977. Freshwater and terrestrial mollusks. Pp.143-194. IN: J.E. Cooper, S.S. Robinson and J.B. Funderburg (eds.) Endangered and threatened plants and animals of North Carolina. North Carolina State Museum of Natural History, Raleigh. Haas, F. 1969. Superfamilia Unionacea. Das Tierreich (Berlin) 88:x + 663 pp. Johnson, R.I.1970. The systematics and zoogeography of the Unionidae (Mollusca: Bivalvia) of the southern Atlantic slope region. Bulletin of the Museum of Comparative Zoology 140:263-450. Kat, P.W. 1983a. Fossil evidence from Fish House clays for the origin and changes in species composition through time of the northern Atlantic Slope unionid fauna (Mollusca: Bivalvia). Proceedings of the Academy of Natural Sciences of Philadelphia 135:85-101. Kat, P.W. 1983b. Patterns of electrophoretic and morphologic variability in a widely distributed unionid: an initial survey. The Netherlands Journal of Zoology 33(1):21-40. Kat, P.W. and G.M. Davis. 1984. Molecular genetics of peripheral populations of Nova Scotian Unionidae (Mollusca: Bivalvia). Biological Journal of the Linnean Society 22:157-185. Lea, I. 1836. A synopsis of the family of Naïades. Philadelphia, Pennsylvania. Carey, Lea and Blanchard, Philadelphia. 1–59, 1 pl. Lea, I. 1836. Description of new fresh water and land shells. Transactions of the American Philosophical Society 6(NS):1-154. Lea, I. 1838. A synopsis of the family of Naïades. 2d ed. enlarged and improved. Carey, Lea and Blanchard, Philadelphia. 44 pp. Lea, I. 1845. Descriptions of new fresh water and land shells. Proceedings of the American Philosophical Society 4:162-168. Lea, I. 1852. Descriptions of new species of the Family Unionidae. Transactions of the American Philosophical Society 10:253-294. Lea, I. 1852. A synopsis of the family of Naïades. 3d ed. greatly enlarged and improved. Blanchard and Lea. Philadelphia, xx + pp. 17–88. Lea, I. 1857. Description of twelve new species of Naiades from North Carolina. Proceedings of the Academy of Natural Sciences 9:85-86. Lea, I. 1860. Descriptions of seven new species of Unionidae from the United States. Proceedings of the Academy of Natural Sciences 12:306-307. Lea, I. 1870. A synopsis of the family Unionidae. 4th ed., very greatly enlarged and improved. Henry C. Lea, Philadelphia, Pennsylvania. xxx + 184 pp. Ortmann, A. E. 1914. Studies in najades (cont.). The Nautilus 28(2):20–22; 28(3):28–34; 28(4):41–47; 28(5[sic]):65–69. Ortmann, A.E. and B. Walker. 1922. On the nomenclature of certain North American naiades. Occasional Papers of the Museum of Zoology, University of Michigan. No. 112:1-75. Simpson, C. T. 1900. Synopsis of the naiades, or pearly fresh-water mussels. Proceedings of the United States National Museum 22(1205):501–1044. Simpson, C. T. 1914. A descriptive catalogue of the naiades, or pearly fresh-water mussels. Parts I–III. Bryant Walker, Detroit, Michigan, xii + 1540 pp. Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 9

Swofford, D. L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Version 4.0b10. Sinauer Associates, Sunderland, Massachusetts. Turgeon, D.D., A.E. Bogan, E.V. Coan, W.K. Emerson, W.G. Lyons, W.L. Pratt, C.F.E. Roper, A. Scheltema, F.G. Thompson, and J.D. Williams. 1988. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 16:vii,1-277,12 color plates Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, M.J. Sweeney, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 26. Second Edition. 536 pp.[Also on CD-ROM]. Wright, S.H. 1897. Contribution to a knowledge of United States Unionidae. The Nautilus 10(12):136-139. Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 10

Table 1 Specimens used in this analysis

Catalog # Locale Drainage N Anodonta cygnea U56842, AF461419 2 Elliptio complanata (topotypes) 5 NCSM 27756 CT: 2 NCSM uncat. Assorted NC locales Cape Fear R. 4 Elliptio congarea NCSM 27922 1 NCSM 27925 1 Elliptio crassidens NCSM uncat. Bear Cr. 1 NCSM uncat. Tombigbee R. 1 NCSN uncat. Alabama R. 1 MMNS 6509 MS: Itawamba Co. E. Fk. Tombigbee R. 1 from Jeanne's stuff 1 Elliptio dilatata NCSM uncat. 1 MMNS 6518 MS: Sunflower Co. Sunflower R. 1 AF156506, AF156507 2 Elliptio fisheriana NCSM 27696 1 NCSM 27697 2 NCSM 27698 1 Elliptio hepaticus from Jeanne's stuff 1 Elliptio icterina NCSM 27919 1 from Jeanne's stuff (A) 1 from Jeanne's stuff (B) 1 from Jeanne's stuff (GA) 1 Elliptio judithae from Jeanne's stuff 1 Elliptio lanceolata NCSM 27729 5 NCSM 27732 5 NCSM 27774 1 from Jeanne's stuff (lanceo01) 1 from Jeanne's stuff (Ulanc1) 1 from Jeanne's stuff (Ulanc2) 1 Elliptio nasutilus NCSM 27691 5 Elliptio producta from Jeanne's stuff (A) 1 from Jeanne's stuff (B) 1 Elliptio roanokensis NCSM 27927 NC: Pee Dee R. 4 Elliptio shephardiana from Jeanne's stuff (A) 1 from Jeanne's stuff (B) 1 Elliptio spinosa from Jeanne's stuff 1 Elliptio steinstansana UAUC 3037 1 Elliptio viridulus from Jeanne's stuff 1 Fusconaia barnesiana NCSM 27692 Duck R. 2 Fusconaia cerina MMNS 6476 MS: Itawamba Co. E. Fk. Tombigbee R. 1 Fusconaia flava MMNS 6516 MS: Sunflower Co. Sunflower R. 1 MMNS 6535 MS: Sunflower Co. Big Sunflower R. 1 Fusconaia masoni from Jeanne's stuff 1 NCSM 27735 VA: 3 NCSM 27736 VA: 3 NCSM 27737 VA: 2 NCSM 27746 VA: Nottoway R. 1 Fusconaia subrotunda NCSM 27783 Hiwassee R. 1 Gonidea angulata AF231755 1 Hyridella menziesi AF231747, AF305369, AF305370 3 Lampsilis cardium AF120653, AF156518, AF156519 3 Lampsilis fasciola AF156520 1 Lampsilis ornata MMNS 6511 MS: Itawamba Co. E. Fk. Tombigbee R. 1 from Jeanne's stuff 1 Lampsilis ovata UAUC 108 1 Lexingtonia dolabelloides Jeanne's stuff 2 Margaritifera auricularia AF303314, AF303315 2 Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 11

Table 1 Continued

Catalog # Locale Drainage N Margaritifera margaritifera AF303340, AF303340 2 Margaritifera margaritifera durrovensis AF303346, AF303347 2 Margaritifera marianae NCSM uncat. 4 Neotrigonia margaritacea U56850 1 Pyganodon cataracta NCSM uncat. NC: Randolph Co. Deep R. 1 Pyganodon grandis MMNS 7157 MS: Sunflower Co. Sunflower R. 1 Pleurobema collina NCSM ???? 1 UAUC 3039 1 UAUC 3040 1 UAUC 3041 1 KKS??? 1 Unio cafferia AF156500, AF156501 2 Unio crassus AF514296 1 Unio pictorum AF468684 1 Unio tumidus AY074807 1

Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 12

Table 2 Pairwise percent sequence divergence (COI) for all specimens of Fusconaia and Lexingtonia analyzed.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 1 F. barnesiana - 2 F. barnesiana 0.00 - 3 F. cerina 8.64 8.64 - 4 F. cerina 8.79 8.79 1.06 - 5 F. flava 9.73 9.73 1.24 1.08 - 6 F. flava 9.31 9.31 1.16 0.97 0.78 - 7 F. flava 9.17 9.17 0.95 0.80 0.64 0.00 - 8 F. flava 8.96 8.96 2.04 1.80 1.59 1.39 1.35 - 9 F. flava 11.82 11.82 12.27 12.73 13.04 12.58 12.55 13.44 - 10 F. flava 9.25 9.25 1.36 1.21 0.94 0.97 0.80 1.78 12.28 - 11 F. flava 8.64 8.64 1.21 1.06 0.94 0.20 0.48 1.13 11.97 1.06 - 12 F. masoni 9.05 9.05 2.77 2.60 2.78 2.14 2.46 3.38 12.27 2.61 2.61 - 13 F. masoni 8.88 8.88 2.92 2.76 2.94 2.14 2.62 3.38 12.10 2.77 2.44 0.16 - 14 F. masoni 9.04 9.04 2.82 2.65 2.84 2.13 2.51 3.38 12.13 2.66 2.32 0.17 0.00 - 15 F. masoni 8.20 8.20 3.31 2.82 3.33 2.91 2.83 3.57 11.78 3.15 3.14 1.00 1.17 1.17 - 16 F. masoni 8.72 8.72 3.74 3.26 3.76 2.72 3.27 3.36 12.39 3.58 3.58 0.83 0.99 1.00 0.17 - 17 F. masoni 8.79 8.79 2.73 2.58 2.78 2.12 2.38 3.35 12.42 2.58 2.42 0.33 0.16 0.00 1.16 1.64 - 18 F. masoni 8.03 8.03 3.18 2.73 3.25 2.69 2.68 3.33 12.12 3.03 3.18 0.81 0.97 0.98 0.16 0.50 1.06 - 19 F. masoni 8.94 8.94 2.58 2.42 2.62 2.12 2.22 3.35 12.58 2.43 2.58 0.16 0.31 0.16 0.99 1.47 0.15 0.91 - 20 F. masoni 9.52 9.52 3.10 2.94 3.11 2.14 2.78 3.38 12.86 2.94 2.94 0.00 0.17 0.17 0.99 0.81 0.67 1.30 0.50 - 21 F. masoni 8.92 8.92 2.71 2.52 2.73 2.33 2.34 3.46 12.39 2.52 2.12 0.19 0.19 0.19 1.16 0.97 0.20 0.97 0.20 0.19 - 22 F. subrotunda 7.27 7.27 3.79 4.24 4.34 4.06 3.97 4.53 11.67 3.94 3.79 3.92 4.08 3.99 4.15 4.57 3.79 3.94 3.64 4.24 3.30 - 23 L. dollabelloides 5.87 5.87 7.49 8.06 8.31 7.74 7.74 7.83 11.78 7.89 7.67 7.94 7.94 7.93 7.35 7.54 7.87 7.48 7.87 7.94 7.93 7.70 - 24 L. dollabelloides 5.70 5.70 7.88 8.07 8.69 8.13 8.13 8.28 11.78 8.27 8.06 8.32 8.32 8.31 7.34 7.53 8.25 7.48 8.25 8.32 8.32 8.08 0.58 -

Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 13

Table 2 Pairwise percent sequence divergence (COI) for all specimens of Elliptio analyzed.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 1 E. complanata CT - 2 E. complanata CT 0.00 - 3 E. complanata NC 2.20 2.20 - 4 E. complanata NC 2.43 2.43 0.16 - 5 E. complanata NC 2.38 2.38 0.16 0.33 - 6 E. complanata NC 2.71 2.71 0.48 0.32 0.64 - 7 E. complanata (TOPO) 0.15 0.15 2.05 2.28 2.23 2.55 - 8 E. complanata (TOPO) 0.15 0.15 2.05 2.28 2.23 2.55 0.00 - 9 E. complanata (TOPO) 0.15 0.15 2.05 2.28 2.23 2.55 0.00 0.00 - 10 E. complanata (TOPO) 0.30 0.30 2.05 2.28 2.23 2.55 0.00 0.00 0.00 - 11 E. complanata (TOPO) 0.15 0.15 2.05 2.28 2.23 2.55 0.00 0.00 0.00 0.00 - 12 E. congarea 2.12 2.12 2.05 2.28 2.24 2.56 1.88 1.88 1.87 1.86 1.89 - 13 E. congarea 1.97 1.97 2.21 2.29 2.39 2.72 1.72 1.72 1.71 1.70 1.73 1.36 - 14 E. crassidens 1.82 1.82 1.58 1.80 1.76 2.08 1.40 1.40 1.40 1.39 1.42 1.21 1.06 - 15 E. crassidens 1.82 1.82 1.42 1.63 1.60 1.92 1.56 1.56 1.56 1.54 1.57 1.21 1.06 0.15 - 16 E. crassidens 1.67 1.67 1.58 1.80 1.76 2.08 1.40 1.40 1.40 1.39 1.42 1.07 0.91 0.31 0.46 - 17 E. crassidens 1.39 1.39 1.44 1.64 1.61 1.93 1.11 1.11 1.10 1.10 1.12 0.94 0.78 0.16 0.31 0.15 - 18 E. crassidens 3.05 3.05 3.47 3.68 3.66 4.06 2.89 2.89 2.88 2.89 2.89 2.90 2.32 1.54 1.72 1.92 1.75 - 19 E. dilatata 8.07 8.07 8.09 7.81 7.98 8.62 7.94 7.94 7.94 7.95 7.93 7.12 7.60 7.44 7.59 7.13 6.94 8.72 - 20 E. dilatata 7.42 7.42 7.24 7.30 7.13 7.77 7.31 7.31 7.30 7.39 7.23 6.67 7.12 7.12 7.13 6.68 6.71 8.08 1.12 - 21 E. dilatata 8.43 8.43 8.22 7.49 8.10 8.74 8.30 8.30 8.30 8.30 8.21 7.49 7.65 7.81 7.96 7.50 7.31 8.34 0.79 1.41 - 22 E. dilatata 8.58 8.58 8.37 7.65 8.26 8.90 8.46 8.46 8.45 8.46 8.36 7.65 7.80 7.96 8.12 7.66 7.47 8.53 0.95 1.57 0.16 - 23 E. fisheriana 6.86 6.86 7.37 7.56 7.55 7.89 6.73 6.73 6.73 6.73 6.71 6.04 6.04 6.03 6.19 6.37 6.23 6.46 8.71 8.19 8.40 8.56 24 E. fisheriana 6.52 6.52 7.03 7.23 7.21 7.55 6.39 6.39 6.39 6.40 6.37 5.69 5.69 5.68 5.84 6.02 5.89 6.25 8.38 7.85 8.06 8.22 25 E. fisheriana 6.68 6.68 7.20 7.40 7.38 7.72 6.56 6.56 6.55 6.56 6.54 5.85 5.86 5.85 6.01 6.19 6.06 6.45 8.55 8.01 8.23 8.39 26 E. fisheriana 6.73 6.73 7.23 7.42 7.41 7.74 6.60 6.60 6.60 6.60 6.58 5.91 6.08 5.91 6.07 6.24 6.11 6.46 8.39 7.89 8.08 8.25 27 E. hepaticus 1.19 1.19 2.22 2.43 2.42 2.82 1.00 1.00 1.00 1.00 1.00 1.82 1.23 1.21 1.40 1.21 1.03 2.07 7.83 7.20 7.45 7.63 28 E. icterina 2.12 2.12 0.79 0.98 0.96 1.29 1.86 1.86 1.86 1.85 1.88 1.82 1.97 1.52 1.52 1.37 1.24 3.25 8.22 7.73 8.57 8.73 29 E. icterina 1.78 1.78 1.99 2.21 2.20 2.60 1.60 1.60 1.59 1.60 1.60 2.39 1.80 1.39 1.58 1.39 1.20 2.42 8.36 7.71 7.97 8.16 30 E. icterina 1.76 1.76 1.94 2.13 2.13 2.52 1.57 1.57 1.57 1.57 1.56 2.57 1.99 1.59 1.77 1.59 1.39 2.77 8.75 8.14 8.37 8.56 31 E. icterina 1.75 1.75 1.95 2.16 2.15 2.54 1.57 1.57 1.57 1.57 1.57 2.56 1.98 1.58 1.77 1.58 1.38 2.77 9.18 8.56 8.80 8.99 32 E. judithae 2.70 2.70 0.96 1.16 1.15 1.54 2.53 2.53 2.52 2.53 2.52 3.12 2.74 2.34 2.14 2.34 2.15 3.35 9.09 8.47 8.71 8.90 33 E. lanceolata 10.53 10.53 10.91 10.80 11.10 11.10 10.42 10.42 10.41 10.41 10.41 9.71 10.21 10.05 10.21 10.05 9.92 11.51 9.59 9.37 9.25 9.41 34 E. lanceolata 9.92 9.92 10.63 10.52 10.82 10.82 9.80 9.80 9.79 9.80 9.79 9.43 9.77 9.77 9.93 9.77 9.64 11.51 9.30 9.09 8.96 9.13 35 E. lanceolata 10.02 10.02 10.73 10.61 10.92 10.92 9.90 9.90 9.90 9.90 9.90 9.54 9.87 9.87 10.03 9.88 9.74 11.51 9.41 9.20 9.07 9.24 36 E. lanceolata 10.28 10.28 11.00 10.89 11.19 11.19 10.17 10.17 10.16 10.16 10.16 9.80 10.30 10.14 10.29 10.14 10.01 11.52 9.70 9.50 9.36 9.53 37 E. lanceolata 10.19 10.19 10.91 10.80 11.10 11.10 10.08 10.08 10.07 10.08 10.08 9.71 10.21 10.05 10.21 10.05 9.92 11.51 9.59 9.37 9.25 9.41 38 E. lanceolata 10.61 10.61 11.33 11.22 11.52 11.52 10.50 10.50 10.49 10.49 10.49 10.13 10.63 10.47 10.63 10.47 10.34 11.73 10.03 9.83 9.69 9.86 39 E. lanceolata 10.19 10.19 10.91 10.80 11.10 11.10 10.08 10.08 10.07 10.08 10.08 9.71 10.21 10.05 10.21 10.05 9.92 11.51 9.59 9.37 9.25 9.41 40 E. lanceolata 10.42 10.42 10.79 10.68 10.98 10.98 10.30 10.30 10.29 10.30 10.29 9.60 9.93 9.93 10.09 9.93 9.80 11.51 9.46 9.25 9.12 9.29 41 E. lanceolata 10.56 10.56 10.93 10.82 11.12 11.13 10.44 10.44 10.43 10.44 10.44 9.73 10.07 10.07 10.23 10.07 9.94 11.53 9.61 9.40 9.27 9.44 42 E. lanceolata 10.22 10.22 10.93 10.82 11.12 11.13 10.10 10.10 10.10 10.10 10.10 9.73 10.07 10.07 10.23 10.07 9.94 11.53 9.61 9.40 9.27 9.44 Table 2 Continued

Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 14

23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 23 E. fisheriana - 24 E. fisheriana 0.84 - 25 E. fisheriana 1.01 0.17 - 26 E. fisheriana 0.17 0.68 0.85 - 27 E. hepaticus 5.71 5.49 5.70 5.70 - 28 E. icterina 7.18 6.83 7.00 7.04 2.02 - 29 E. icterina 6.65 6.43 6.63 6.65 1.04 1.78 - 30 E. icterina 6.50 6.28 6.48 6.49 0.81 1.76 0.81 - 31 E. icterina 6.90 6.69 6.88 6.90 1.44 1.76 1.41 1.18 - 32 E. judithae 7.05 6.84 7.04 7.05 2.47 1.54 2.05 1.98 1.79 - 33 E. lanceolata 9.77 9.10 9.27 9.62 10.15 10.35 10.64 11.20 11.24 11.91 - 34 E. lanceolata 9.66 8.99 9.16 9.50 10.16 10.07 10.65 11.20 11.24 11.90 0.51 - 35 E. lanceolata 9.80 9.13 9.30 9.60 10.14 10.18 10.63 11.19 11.23 11.91 0.34 0.17 - 36 E. lanceolata 9.77 9.10 9.27 9.72 10.15 10.44 10.65 11.20 11.24 11.92 0.34 0.17 0.00 - 37 E. lanceolata 9.77 9.10 9.27 9.62 10.16 10.35 10.64 11.21 11.24 11.91 0.51 0.34 0.17 0.17 - 38 E. lanceolata 9.96 9.29 9.46 10.05 10.38 10.78 10.86 11.42 11.46 12.13 0.51 0.34 0.17 0.17 0.34 - 39 E. lanceolata 9.77 9.10 9.27 9.62 10.15 10.35 10.64 11.20 11.24 11.91 0.34 0.17 0.00 0.00 0.17 0.17 - 40 E. lanceolata 9.82 9.14 9.32 9.66 9.97 10.24 10.46 11.02 11.06 11.91 0.51 0.68 0.51 0.51 0.68 0.68 0.51 - 41 E. lanceolata 9.79 9.12 9.29 9.64 9.99 10.38 10.48 11.04 11.08 11.93 0.51 0.68 0.51 0.51 0.68 0.68 0.51 0.00 - 42 E. lanceolata 9.79 9.12 9.29 9.64 10.18 10.38 10.66 11.23 11.26 11.94 0.34 0.17 0.00 0.00 0.17 0.17 0.00 0.51 0.51 - 43 E. lanceolata 9.49 8.82 8.99 9.51 9.48 10.15 9.76 10.35 10.39 11.24 1.70 1.89 1.70 1.69 1.86 2.03 1.70 1.54 1.54 1.71 - 44 E. lanceolata 9.88 9.28 9.47 9.89 9.73 10.51 10.23 10.79 10.83 11.68 1.35 1.54 1.35 1.35 1.54 1.55 1.35 1.15 1.15 1.35 1.54 - 45 E. lanceolata 9.88 9.28 9.47 9.89 9.73 10.51 10.23 10.79 10.83 11.68 1.35 1.54 1.35 1.35 1.54 1.55 1.35 1.15 1.15 1.35 1.54 0.39 - 46 E. lanceolata 10.07 9.47 9.66 10.08 10.11 10.73 10.60 11.15 11.20 11.88 0.38 0.19 0.00 0.00 0.19 0.19 0.00 0.58 0.58 0.00 1.75 1.35 1.35 - 47 E. nasutilus 1.67 0.83 0.99 1.49 5.86 7.58 6.78 6.65 7.05 7.19 9.54 9.42 9.53 9.66 9.54 9.99 9.54 9.59 9.57 9.57 9.85 9.60 9.60 9.82 48 E. nasutilus 1.67 0.83 0.99 1.49 5.86 7.58 6.78 6.65 7.05 7.19 9.54 9.42 9.53 9.66 9.54 9.99 9.54 9.59 9.57 9.57 9.85 9.60 9.60 9.82 49 E. nasutilus 1.67 0.83 0.99 1.49 5.86 7.58 6.78 6.65 7.05 7.19 9.54 9.42 9.53 9.66 9.54 9.99 9.54 9.59 9.57 9.57 9.85 9.60 9.60 9.82 50 E. nasutilus 1.67 0.83 0.99 1.49 5.86 7.58 6.78 6.65 7.05 7.19 9.54 9.42 9.53 9.66 9.54 9.99 9.54 9.59 9.57 9.57 9.85 9.60 9.60 9.82 51 E. nasutilus 1.67 0.83 0.99 1.49 5.86 7.58 6.78 6.65 7.05 7.19 9.54 9.42 9.53 9.66 9.54 9.99 9.54 9.59 9.57 9.57 9.85 9.60 9.60 9.82 52 E. producta 5.36 5.55 5.56 5.36 4.66 5.42 4.91 4.25 4.91 5.13 9.90 9.91 9.89 9.90 9.90 10.13 9.90 9.71 9.73 9.92 8.99 9.47 9.47 9.86 53 E. producta 5.82 6.00 6.20 5.82 4.71 5.33 4.82 4.41 5.04 5.46 9.80 9.81 9.79 9.80 9.80 10.02 9.80 9.62 9.64 9.82 8.93 9.38 9.38 9.76 54 E. roanokensis 7.35 7.01 7.17 7.07 2.23 1.06 1.99 1.97 1.97 0.97 10.85 10.57 10.67 10.96 10.85 11.30 10.85 10.74 10.87 10.87 10.46 11.09 11.09 11.30 55 E. roanokensis 7.36 7.01 7.18 7.22 2.23 0.91 2.00 1.98 1.97 0.97 10.87 10.58 10.69 10.95 10.87 11.29 10.87 10.75 10.89 10.89 10.61 10.91 10.91 11.13 56 E. roanokensis 7.35 7.00 7.17 7.21 2.82 1.21 2.58 2.54 2.54 1.55 10.19 9.91 10.02 10.28 10.19 10.62 10.19 10.07 10.21 10.21 10.00 10.33 10.33 10.54 57 E. roanokensis 7.18 6.83 7.00 7.04 2.02 0.61 1.78 1.76 1.76 0.77 10.69 10.40 10.51 10.77 10.69 11.11 10.69 10.57 10.71 10.71 10.46 10.90 10.90 11.11 58 E. shephardiana 6.03 6.20 6.40 6.02 5.09 5.73 5.19 5.28 5.69 5.85 10.27 10.26 10.26 10.27 10.27 10.48 10.27 10.26 10.28 10.29 9.62 10.04 10.04 10.23 59 E. shephardiana 6.03 6.20 6.40 6.02 5.09 5.73 5.19 5.28 5.69 5.85 10.27 10.26 10.26 10.27 10.27 10.48 10.27 10.26 10.28 10.29 9.62 10.04 10.04 10.23 60 E. spinosa 7.37 7.16 7.36 7.37 6.91 8.05 7.42 7.42 7.86 8.58 12.01 12.01 12.01 12.02 12.02 12.23 12.01 12.01 12.03 12.04 11.55 11.97 11.97 11.97 61 E. steinstansana 9.11 8.51 8.71 9.12 8.10 7.42 7.58 8.00 8.03 8.77 9.86 9.86 9.87 9.88 9.87 10.08 9.86 9.87 9.88 9.89 8.64 9.65 9.27 9.85 62 E. viridulus 0.58 0.00 0.20 0.58 5.46 6.48 6.39 6.25 6.66 6.82 9.50 9.49 9.49 9.51 9.51 9.72 9.50 9.50 9.52 9.53 9.03 9.27 9.27 9.46 Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 15

Table 2 Continued

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 43 E. lanceolata 10.46 10.46 10.70 10.37 10.78 10.79 10.57 10.57 10.56 10.62 10.68 9.70 10.30 10.00 10.01 9.87 9.67 10.82 9.52 9.24 9.84 10.00 44 E. lanceolata 10.89 10.89 11.16 11.03 11.37 11.37 10.77 10.77 10.76 10.77 10.77 9.78 10.17 10.17 10.35 10.17 10.03 11.28 9.81 9.56 9.42 9.61 45 E. lanceolata 10.89 10.89 11.16 11.03 11.37 11.37 10.77 10.77 10.76 10.77 10.77 9.78 9.98 10.17 10.35 10.17 10.03 11.28 9.42 9.18 9.04 9.22 46 E. lanceolata 10.73 10.73 11.36 11.22 11.57 11.57 10.59 10.59 10.59 10.59 10.59 10.00 10.38 10.38 10.56 10.38 10.23 11.48 10.01 9.78 9.62 9.81 47 E. nasutilus 7.27 7.27 7.56 7.79 7.77 8.10 6.84 6.84 6.83 6.93 6.91 6.52 6.82 6.97 6.98 6.83 6.57 6.77 8.55 8.18 8.58 8.74 48 E. nasutilus 7.27 7.27 7.56 7.79 7.77 8.10 6.84 6.84 6.83 6.93 6.91 6.52 6.82 6.97 6.98 6.83 6.57 6.77 8.55 8.18 8.58 8.74 49 E. nasutilus 7.27 7.27 7.56 7.79 7.77 8.10 6.84 6.84 6.83 6.93 6.91 6.52 6.82 6.97 6.98 6.83 6.57 6.77 8.55 8.18 8.58 8.74 50 E. nasutilus 7.27 7.27 7.56 7.79 7.77 8.10 6.84 6.84 6.83 6.93 6.91 6.52 6.82 6.97 6.98 6.83 6.57 6.77 8.55 8.18 8.58 8.74 51 E. nasutilus 7.27 7.27 7.56 7.79 7.77 8.10 6.84 6.84 6.83 6.93 6.91 6.52 6.82 6.97 6.98 6.83 6.57 6.77 8.55 8.18 8.58 8.74 52 E. producta 4.99 4.99 5.24 5.49 5.46 5.88 4.82 4.82 4.82 4.82 4.82 4.62 4.62 3.76 3.95 4.18 4.01 4.49 7.76 7.52 7.36 7.56 53 E. producta 4.93 4.93 5.16 5.39 5.37 5.77 4.77 4.77 4.76 4.77 4.76 4.55 4.56 3.75 3.94 4.15 3.99 4.63 7.92 7.67 7.53 7.72 54 E. roanokensis 2.27 2.27 0.48 0.67 0.65 0.98 2.18 2.18 2.18 2.31 2.21 2.27 2.42 1.97 1.67 1.83 1.71 3.46 7.92 7.27 8.28 8.43 55 E. roanokensis 2.42 2.42 0.48 0.67 0.65 0.98 2.18 2.18 2.17 2.16 2.20 1.97 2.27 1.82 1.52 1.67 1.56 3.46 8.07 7.58 8.42 8.58 56 E. roanokensis 2.73 2.73 0.79 0.66 0.96 0.97 2.48 2.48 2.48 2.46 2.51 2.27 2.58 2.12 1.82 1.98 1.86 4.03 8.07 7.58 8.42 8.58 57 E. roanokensis 2.12 2.12 0.16 0.34 0.33 0.65 1.86 1.86 1.86 1.85 1.88 1.82 1.97 1.52 1.21 1.37 1.24 3.25 7.91 7.42 8.26 8.42 58 E. shephardiana 5.52 5.52 5.56 5.77 5.75 6.14 5.37 5.37 5.37 5.37 5.37 4.81 4.81 4.03 4.21 4.41 4.25 5.06 8.06 7.83 7.68 7.87 59 E. shephardiana 5.52 5.52 5.56 5.77 5.75 6.14 5.37 5.37 5.37 5.37 5.37 4.81 4.81 4.03 4.21 4.41 4.25 5.06 8.06 7.83 7.68 7.87 60 E. spinosa 7.26 7.26 8.27 8.50 8.47 8.86 7.11 7.11 7.11 7.11 7.11 6.36 6.92 6.53 6.71 6.92 6.76 7.21 10.19 9.96 9.82 10.00 61 E. steinstansana 8.55 8.55 8.25 8.12 8.46 8.46 8.43 8.43 8.42 8.43 8.43 7.83 8.04 7.84 8.01 7.84 7.70 8.94 8.82 8.55 8.43 8.23 62 E. viridulus 6.30 6.30 6.71 6.94 6.91 7.30 6.16 6.16 6.15 6.16 6.14 5.19 5.19 5.18 5.36 5.57 5.42 6.22 8.44 7.82 8.07 8.25

47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 47 E. nasutilus - 48 E. nasutilus 0.00 - 49 E. nasutilus 0.00 0.00 - 50 E. nasutilus 0.00 0.00 0.00 - 51 E. nasutilus 0.00 0.00 0.00 0.00 - 52 E. producta 6.33 6.33 6.33 6.33 6.33 - 53 E. producta 6.75 6.75 6.75 6.75 6.75 0.00 - 54 E. roanokensis 7.73 7.73 7.73 7.73 7.73 5.22 5.14 - 55 E. roanokensis 7.73 7.73 7.73 7.73 7.73 5.02 4.95 0.76 - 56 E. roanokensis 7.73 7.73 7.73 7.73 7.73 4.80 4.74 1.06 0.61 - 57 E. roanokensis 7.58 7.58 7.58 7.58 7.58 5.00 4.93 0.46 0.30 0.61 - 58 E. shephardiana 6.94 6.94 6.94 6.94 6.94 0.42 0.40 5.55 5.37 5.16 5.34 - 59 E. shephardiana 6.94 6.94 6.94 6.94 6.94 0.42 0.40 5.55 5.37 5.16 5.34 0.00 - 60 E. spinosa 7.91 7.91 7.91 7.91 7.91 6.79 6.59 8.25 8.07 8.25 8.05 7.14 7.14 - 61 E. steinstansana 8.63 8.63 8.63 8.63 8.63 7.96 7.75 8.00 8.01 7.81 7.99 8.30 8.30 10.04 - 62 E. viridulus 0.77 0.77 0.77 0.77 0.77 5.53 5.97 6.69 6.51 6.68 6.48 6.18 6.18 7.14 8.49 -

Bogan et al. 2003 Relationships of the Atlantic Pigtoe, Fusconaia masoni 17