ill latin o Museum of

oology

The Systematics and Zoogeography of the

Unionidae (: ) of the Southern Atlantic Slope Region

RICHARD I. JOHNSON

HARVARD UNIVERSITY VOLUME 140, NUMBER 6 CAMBRIDGE, MASSACHUSETTS, U.S.A. 27 NOVEMBER 1970 PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY

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© The President and Fellows of Harvard College 1970. THE SYSTEMATICS AND ZOOGEOGRAPHY OF THE (MOLLUSCA: BIVALVIA) OF THE SOUTHERN ATLANTIC SLOPE REGION

RICHARD I. JOHNSON

CONTENTS Apalachieolan and Atlantic Slope re- gions to their unionid faunas .... 283 Introduction 264 Evidence of stream capture .. . 284 266 Acknowledgments The Piedmont Plateau . 285

Part I. The zoogeography of the Unionacea of The Coastal Plain _____ . 286 the Apalachieolan and Atlantic Slope 6. and Conclusions 291 regions 267 Chapter Summary Part II. A revision of the Unionidae from the 1. The distribution of the Union- Chapter St. Marys River, Florida, to the Po- acea in the region as Apalachieolan tomac River, Maryland 293 evidence of a former confluence of the headwaters of the - Introduction 293 Coosa, Apalachicola, and Savannah History of the taxonomists and the collectors river 267 systems of the Atlantic Slope Unionacea 293 defined 267 The Apalachieolan region The taxonomists 293 Analysis of the distribution of the 268 The collectors . _ 294

Chapter 2. The distribution of the Union- The primary systematic studies of Apala- idae in the Southern Atlantic Slope chieolan and Atlantic Slope Union- region as evidence of a former stream acea 294 confluence of the headwaters of the Classification of the Unionacea _ 295 Apalachicola and Savannah river sys- tems 273 Systematic Section 296 The Atlantic defined and Slope region Key to the Unionidae found between the St. subdivided 273 Marys River, Florida, and the Po- Analysis of the distribution of the species 275 tomac River, Maryland 297

PLEUROBEMA - 299 Chapter 3. The Unionacea of the Northern Atlantic Slope region, with a discus- Lexingtonia 300 sion of the probable origin of some Pleurobema (Lexingtonia) collina (Con-

- 300 of the Atlantic slope species . 278 rad) Analysis of the distribution of the species 278 Pleurobema (Lexingtonia) masoni (Con- rad) - 301 Chapter 4. The generic affinities of the 303 Unionacea of the Apalachieolan ELLIPTIC- region, Peninsular Florida, and Canthyria 303 Southern and Northern Atlantic (Canthyria) spinosa (Lea) . - 303 281 Slope regions Elliptio s. s. 304 crassidens crassidens Chapter 5. The relationship of the geo- Elliptio (Elliptio) 305 morphology and topography of the (Lamarck)

: 263 Bull. Mus. Comp. Zool., 140(6) 263-450, November, 1970 264 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Elliptio (Elliptic) crassidens downiei Pyganodon 356 (Lea) - 307 Anodonta (Pyganodon) cataracta cata- Elliptio (Elliptio) congaraea (Lea) ... 308 racta Say 356 310 Elliptio (Elliptio) dariensis (Lea) Anodonta ( Pyganodon ) gibbosa Say .. 359 Elliptio (Elliptio) fraterna (Lea) 312 Anodonta (Pyganodon) implicata Say 360 Elliptio (Elliptio) waccamawensis (Lea) 313 Utterbackia 362 Elliptio (Elliptio) complanata (Light- Anodonta (Utterbackia) imbecilis Say 362 foot) . 314 Anodonta (Utterbackia) couperiana Lea 365 Elliptio (Elliptio) (Lea) 324 hopetonensis STROPHITUS 366 .. Elliptio ( Elliptio ) icterina ( Conrad ) 325 Strophitus undulatus (Say) 367 Elliptio (Elliptio) arctata (Conrad) .. 331 CARUNCULINA 369 Elliptio (Elliptio) lanceolata (Lea) .. 333

338 Caruncidina ( Conrad ) 370 Elliptio ( Elliptio ) shepardiana ( Lea ) pulla UNIOMERUS 339 VILLOSA 371

Uniomerus tetralasmus (Say) 339 Villosa villosa (Wright) . 372 LASMIGONA 343 Villosa vibex (Conrad) 373 Villosa delumbis 375 Platynaias 343 (Conrad) Lasmigona (Platynaias) subviridis (Con- Villosa constricta (Conrad) . 378 rad) ..... 343 LIGUMIA 380 ALASMIDONTA 346 Ligumia nasuta (Say) 380

Prolasmidonta 346 . 382 Alasmidonta heterodon (Prolasmidonta) Lampsilis s. s. 382 (Lea) . 347 Lampsilis (Lampsilis) cariosa (Say) .. 382 Alasmidonta s. s. 348 Lampsilis (Lampsilis) dolabracformis Alasmidonta undulata (Alasmidonta) (Lea) 384 (Say) . 349 Lampsilis (Lampsilis) ovata (Say) .... 386 Alasmidonta (Alasmidonta) triangulata Lampsilis (Lampsilis) ochracea (Say) 388 (Lea) . 351 Lampsilis (Lampsilis) radiata radiata Alasmidonta (Alasmidonta) arcula (Lea) 352 (Gmelin) . 390 Decuramhis 353 Lampsilis (Lampsilis) splendida (Lea) 393 Alasmidonta (Decuramhis) varicosa Bibliography 395 ( Lamarck ) 354

ANODONTA 356 Index to relevant taxa 399

ABSTRACT is also an additional fauna originally of Interior Rasin but whose The Atlantic Slope region consists of origin, ancestry is more remote. those streams flowing into the Atlantic- Ocean from the Altamaha River system, INTRODUCTION , to the lower St. Lawrence River system, Canada, including rivers in New- The Unionacea and Mutelacea (Parodiz foundland and Labrador. Forty species and Ronetto, 1963), or freshwater mussels, comprise the unionid fauna. Most of the are found throughout the world, but it is species that are clearly of Interior Rasin in the Mississippi River system that the origin are a northern group that migrated Unionidae have especially radiated and around the northern end of the Appala- achieved their greatest diversity. This, the chian Mountains before the Pleistocene; Interior Basin, embraces 1,200,000 square there is a southern group that entered the miles. The rivers are very old, and flow Atlantic Slope region through a confluence over vast limestone beds. Here is found a of the Apalachicola and Savannah river variety of shell forms which is rivaled only systems, also in pre-Pleistocene time; there in a few species found in the rivers of Atlantic Slope Unionidae • Johnson 265

China. Within the Interior Basin, H. and species are smaller and are neutral in color. A. van der Schalie (1950: 450) recognized Yet, the only two species of Unionidae with the Ozark and Cumberland each true regions, spines. Elliptic- spinosa (Lea) and of which has an indigenous unionid fauna Pleurobema collina (Conrad), occur here. of its own, as well as the Interior Basin one. Because the Atlantic Slope region has Continental North America is made up distinct northern and southern assemblages of the following additional regions. They of species, it has been divided here into the are given essentially as defined by H. and Southern Atlantic Slope region, which ex- A. van der Schalie (1950). The Pacific tends from the Altamaha River system, region is the area west of the Rocky Georgia, to the James River system, Vir- Mountains. It has a very limited unionid ginia, and the Northern Atlantic Slope fauna, which is clearly of Asiatic origin. region, which extends from the York River Found there are several species of Ano- system, , to the lower St. Lawrence donta, Margaritifera margaritifera (Lin- River system, Canada. Between the York and "Gonidea" River naeus), angulata (Lea), and the glacial drift border ( roughly which may belong to the Asiatic genus along the Pennsylvania- bound- Solenaia Conrad. ary) are found all of the Unionacea that The West Gidf Coastal region consists repopulated the Northern Atlantic Slope of the streams flowing into the Gulf of region north of the limit of glacial drift. Mexico from the eastern slope of Mexico The Unionacea, or freshwater mussels, north to, but not including, the Alabama- offer two advantages for zoogeographic Coosa River system. Found there are study: 1) As shown in the systematic por- several endemic genera of Unionidae. It tion of this paper, there are a reasonable has been tentatively suggested by both number of species, most of which are Simpson (1892: 406) and H. and A. van clearly distinguishable, and whose generic der Schalie (1950: 452) that this might be affinities have been revealed rather clearlv regarded as a subregion of the Interior by Ortmann (1911; 1912a). 2) They have Basin. a limited mode of distribution, being un- The Apalachicolan (or West Floridian) able to pass over land from one drainage region was mentioned by H. and A. van system to another. Their ability to move der Schalie (1950: 450) and is now defined between drainage systems is dependant on as consisting of those river systems flowing the mobility of fishes to which the glochidia into the Gulf of Mexico from the Escambia attach themselves. Of the primary fresh- to the Suwannee, and also including the St. water fishes, Myers (1938: 343) stated, Marys and the Satilla, which flow directly "They are inescapably confined to their own into the Atlantic Ocean. particular drainage systems and can migrate Peninsular Florida has representatives of from one isolated stream basin to the next the slow only six genera. These consist of species only through physiographical change of the land itself (stream capture, etc. [base- derived mostly from the Apalachicolan and leveling] )." Atlantic Slope regions. This area will be of the discussed as a separate region in a sub- For this reason the distribution sequent report. Unionacea gives evidence of former stream The Atlantic Slope region includes the confluences and of flooding in baseleveled river systems flowing into the Atlantic- coastal regions. Ocean from the Altamaha, Georgia, to the Van der Schalie (1945) convincingly lower St. Lawrence, Canada, as well as illustrated the use of Unionidae as a means those of Newfoundland and Labrador. The of tracing major stream confluences. He Unionidae of this region are of Interior also reviewed the old controversy over Basin origin, but on the Atlantic slope the mechanical distribution, which implies that 266 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

unionid distribution is haphazard and for- ception of Margaritifera mar^aritifera caused birds to whose Linnaeus Alasmidonta tuitous, by aquatic ( ) , marginata ( Say ) , feet gravid females may occasionally be- and Anodonta catorocta fragilis (Lamarck), come attached. This theory postulates that which are beyond the scope of this paper, the shell will be carried to a different river all of the Atlantic Slope Unionacea are system, and that there the glochidia will included and their nomenclature is revised. be released and will find a suitable host fish. No one has actually established that ACKNOWLEDGMENTS Unionidae have been successfully spread In Dr. 1961, William J. Clench, my long- by birds in this manner, but even granting time mentor, received a grant from the this there is no evidence that possibility, National Science Foundation (G18922) to it is, or was, an method of dis- important make a collection of the freshwater tribution, since zoogeographic data fail to mollusks of the Atlantic Slope of Georgia. substantiate unionid distribution by any Upon completion of the expedition, he agent other than fish. generously gave me the Unionidae for In their discussion of the freshwater mol- study. As the study progressed, it gradu- lusks of the Apalachicolan region, Clench ally became apparent that the unionids of and Turner (1956: 103) support the idea the entire Southern Atlantic slope had to that that fauna was distributed by me- be included if the work was to be of any chanical means, though they are vague as significance. to what those means might have been. The opportunity for examining relevant Exception is taken here to their theory of types was provided by: Dr. Norman distribution, but only as far as the Rritish Museum Natural Tebble, ; Unionacea are concerned. ( History ) Dr. Gilbert Ranson, Paris Museum; Dr. The Unionacea were re- Apalachicolan G. Mermod, Geneva Museum; Dr. Arthur studied, and it is demonstrated here that H. Clarke, Jr., National Museum of Can- their general distribution is not fortuitous. ada; Dr. David H. Stansbery, State References are made to some of the Museum; Dr. Juan Parodiz, Carnegie Mu- Unionidae of the Alabama-Coosa River seum, Pittsburgh; Dr. R. Tucker Abbott, system to illustrate the of some of origin Academy of Natural Sciences of Philadel- the Apalachicolan species and to illuminate phia; Dr. Henry van der Schalie, Museum the sequence of stream captures which of Zoology, University of . Drs. affected the distribution of both Apalachi- Harold Render, Joseph Rosewater, and colan and Southern Atlantic Slope Unioni- National dae. Joseph Morrison, Museum, me to the In his classic study of the influence of kindly permitted study collections under their care and to borrow the Alleghenian Divide on the distribution of all relevant types. The cost of photo- mollusks and crayfish, Ortmann ( 1913a ) assumed that the Unionidae were dis- graphing the types was generously borne by tributed by natural means. Roth by choice the William F. Milton Fund, Harvard Uni- and because of the confusion in which he versity. found the systematics of the southern Dr. Ruth Patrick, Academy of Natural species, his work was limited to the Sciences of Philadelphia, made available Atlantic side of the divide, primarily to the Unionidae collected for her by John the species of Unionacea found on the N. Rates and Samuel L. H. Fuller; Mr. Northern Atlantic slope. Rates allowed me to study the portion of In the present study, the distribution of this material under his care at the Museum the Apalachicolan and Atlantic Slope of Zoology, University of Michigan. Dr. Unionacea is analyzed, and, with the ex- Patrick kindly deposited most of the Atlantic Slope Unionidae • Johnson 267

material collected by Mr. Fuller in the William II. Heard, University of Florida. Museum of Comparative Zoology. Tallahassee, for specimens. Herbert D. Athearn, Cleveland, Ten- Finally, thanks are offered to Drs. Ken- is for the neth nessee, gratefully acknowledged J. Boss, Raymond A. Paynter, Jr., and material which he has presented to the Ruth D. Turner, who critically read the Museum of Comparative Zoology over the manuscript, and to Drs. Arthur II. Clarke. years, and which has proved useful in this Jr. and Joseph P. E. Morrison, who offered study. Thanks are also extended to Dr. opinions on some matters of .

PART I. THE ZOOGEOGRAPHY OF THE UNIONACEA OF THE APALACHICOLAN AND ATLANTIC SLOPE REGIONS

in CHAPTER 1 cola river systems are absent the Savannah River system. The distribution of the Unionacea in the To understand the origins of the Unioni- Apalachicolan region as evidence of a dae of the Southern Atlantic Slope, it was former confluence of the headwaters of the necessary to study the distribution of the Alabama-Coosa, Apalachicola, and Savan- superfamily Unionacea in the Alabama- nah river systems. Coosa and Apalachicola river systems. Figure 1 illustrates the propinquity of Further, the whole Apalachicolan region the headwaters of the Alabama-Coosa, had to be considered, to determine if it Apalachicola, and Savannah river systems. could be established that unionid distri- Van der Schalie (1945) made it clear that bution there is not fortuitous. there was once a connection between the Tfie Apalachicolan region defined. This Tennessee and Alabama river systems; this region is regarded here as consisting of the connection is of interest here since certain river systems flowing into the Gulf of unionid species of the Alabama-Coosa Mexico, from the Escambia to the Su- River system are discussed in this paper. wannee. Also included are the St. Marys in Matteson (1948a: 131) following, part, and Satilla, although they flow into the and (1894), suggested 1 as Table Hayes Campbell Atlantic Ocean ( Plate ) , because, that in the late the Chattahoochee Tertiary, 1 shows, their modest unionid faunas con- River of the River system Apalachicola sist entirely of species found in the Apa- captured a tributary of the Etowah River, lachicolan region, and further, because of the Alabama-Coosa River system, and EUiptio crassidens crassidens and E. c. that the Savannah River then captured one downiei occur in them, respectively, and of the tributaries of the Chattahoochee are dominant. River. This present study supports these According to Cooke (1945: 273), in the geomorphological assertions, but in a dif- early Pleistocene, during the Aftonian ferent order. The distribution of Apalachi- interglacial stage when the Brandywine colan species in the Savannah River system terrace Citronelle Formation in the South- seems to indicate that this system may ( was formed, the sea level was 270 first have been connected to, and later east) feet above the present level. If this were separated from, the Chattahoochee River of the area the before the latter was connected to the so, most occupied by Alabama-Coosa River. These conditions present St. Marys and Satilla river systems, and all of Peninsular Florida, would explain why certain species found virtually of in both the Alabama-Coosa and Apalachi- were inundated, with the exception 268 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Text-figure 1. Northern Georgia, illustrating the propinquity of the headwaters of the Etowah River (ER), of the Alabama-Coosa; of the Chattahoochee River (CR), of the Apalachicola; and of the Savannah (HSR) river systems. The topographic areas are the Appalachian Mountains (AM), Appalachian Valley (AV), and the Piedmont Plateau (PP). The Blue Ridge divide (BR) is indicated by large dots. The minor divides are indicated by smaller dots. (After Keith, 1925, pl. 29.)

several small islands in the vicinity of ida was reduced to a number of small is- present Polk County (Cooke, 1945: 273). x lands where a unionid fauna might have It is now generally agreed that this flood- had refugia. In any case, except for a small ing took place in the Upper Miocene (Alt portion of the Satilla River which is above and Brooks, 1965: 408). the area of maximum flooding, the remain- MacNeil (1950: 98, 99) casts doubt on der of it, and all of the St. Marys River, is Cooke's evidence on the extent of Aftonian of more recent origin (see p. 289). flooding, and on the basis of his identifi- It is assumed that these two rivers were cation of marine terraces that the concludes mostly repopulated from the west. The ap- Citronelle Formation is of subaerial origin, pearance of Elliptio c. cra.ssiden.s in the and that there is no evidence to indicate Pliocene of Florida (see note on p. 271) that the sea ever it to an alti- transgressed indicates that it has been present in this tude of more than 150 feet the during general area over a long period of time. Yarmouth interglacial stage. This flooding is now thought to have occurred during the Analysis of the distribution of the species. Pliocene. MacNeil's detailed map (pl. 19) Atlantic shows that even at 150 feet Peninsular Flor- The Apalachicolan and Slope regions consist of a number of independent 1 Orange (Ocala) Island, referred to by Clench river systems, some with quite different and Turner was a land that (1950: 104), mass faunas. Table 1 shows all of the Unionacea was separated from the continent by the Suwannee known from the individual Strait dining the late Oligocene (Vanghan, 1910: Apalachicolan 156), and its existence appears to have had no river systems, systematically arranged. bearing on the present nnionid fauna. Some differences in species concepts and Atlantic Slope Unionidae • Johnson 269

Table 1. Distribution ok the Umonacea of the Apalachk :olan hegion and RELEVANT SPECIES OF THE AlABAMA-CooSA RlYKR SYMHM. 270 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Table 1. (Continued)

c - c S 8 0) tb O a, -^ o IS ^ rt t/i C w W5 «

* -" is 5 — ii 1) > > -S.S; a O w« >-« u5 Ok

46. Lampsilis subangulata (Lea) X X X 47. Medionidus penicillatus (Lea) X 48. Medionidus walkeri (Wright) (7) X X X X 49. Glebula rotundata (Lamarck) X X TOTAL NUMBER OF SPECIES 23 11 18 33 20 13 4

* (See Johnson, 1965; 1967b, c; 1968; 1969a). A. perplicata (Conrad) has not been previously published on as part of the Apalachicolan fauna. It has been found in the Escambia River, Pine Barren, Escambia Co., Florida, and in Picket Retch Lake [not located, but presumed to be in the Yellow River system], Walton Co., Florida. 1. Fusconaia, represented by several species. 2. Pleurobcma rubellum (Lea).

3. Elliptic) dilatata ( Rafinesque) see p. 279. 4. Alasmidonta mccordi Athearn (1964: 134, pi. 9, figs. a-b). Not close to Alasmidonta s.s. 5. unicolor (Lea). of 6. Clench and Turner (1956: 158), following Simpson (1914: 91), recognize floridensis ( Lea ) as a subspecies of the form would be Lampsilis anodontoides ( Lea )= teres (Rafinesque), but admit that "young specimens typical difficult to separate from this subspecies." It is true that specimens )f this species tend to be smaller and thinner toward the southern end of its range, but floridensis is not a subspecies as this concept is currently understood. It was de scribed as: Unio floridensis Lea 1852. Trans. Amer. Philos. Soc. 10: 274, pi. 21, fig. 31 [Chactahachi River, West Florida, figured holotvpe AXSP 42081. Clench and Turner (1956: 202) restricted the tvpe locality to the Choctawhatchee River, 1 mi. W Caryville, Holmes Co., Florida]. Lea, 1852, Obs. Unio, 5: 30. 7. Clench and Turner (1956: 189, pi. 6, fig. 7), under the name Medionidus penicillatus (Lea), described and figured Medionidus walkeri (Wright), a species endemic to the Apalachicolan region having a well defined posterior ridge and a posterior slope with strong corrugations that run from the posterior ridge to the posterior margin. Medionidus penicillatus (Lea), though found in the Apalachicolan region, is more common in the Alabama-Coosa River system. It has a very rounded posterior ridge, and a posterior slope that has, when sculpture is present, small, slightly curved folds that run from the posterior ridge to the posterior margin. It was described as: Unio penicillatus Lea 1857, Proc. Acad. Nat Sci. Phila., 9: 171 (Chattahoochee River, near Columbus [Muscogee Co.]; [Chattahoochee River], near Atlanta [De Kalb Co.]; Flint River near Albany, [Dougherty Co.]; all Georgia). Lea, 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 203, pi. 23, fig 85, figured holotvpe USNM 84142 from the Flint River. Clench and Turner (1956: 190) restricted the type-locality to the Chattahoochee River, near Columbus). Lea, 1859, Obs. Unio., 7: 21. nomenclature from that of Clench and others have spread in the opposite direc- Turner (1956) were noted by Johnson tion. (1965; 1967b, 1967c; 196S; 1969a); the re- Six Apalachicolan unionids are so ubiq- in uitous the mainder are discussed the notes to Table throughout region ( Table 1, 1, or in the systematic portion of this paper. A) that they offer no clues as to their Because of its former connection with method of distribution. Amblema perpli- the , the unionid fauna of cata (Conrad) is found in rivers flowing the Alabama-Coosa River system is ex- into the Gulf of Mexico from central Texas tensive. Only those genera and species to the Yellow River system, Florida, and that seem germane to the understanding in the west, north to river systems in Arkan- of the distribution of the Unionacea of the sas and Missouri. It is close to A. plicata Apalachicolan and Atlantic Slope regions (Say), which is widely distributed through- are included here. out the Interior Basin. Lampsilis exca- Forty-nine species comprise the unionid vatus ranges uniformly from the Pearl fauna of the Apalachicolan region (Table River, Mississippi, to the Escambia River 1 nineteen affinities Florida. a ) ; species have with system, Margaritifera hembeli, species to the west, twelve are endemic to member of the family Margaritiferidae, the region, and eleven more are restricted appears to be a relict. It is found in one to individual river systems. Three species, other drainage, the Bayou Teehe, Louisi- which were probably originally endemic ana, in the West Gulf Coastal region. Ano- to the Apalachicolan region, have spread donta suborbiculata is widely distributed into the Atlantic Slope region, and four throughout the Interior Basin. Atlantic Slope Unionidae • Johnson 271

Table 1, A. Unionacea, derived from the west, found in the Ai.abama-Coosa River system, and which terminate in the Apalachk :<>i.a\ region, or are widely distributed throughout and beyond it.

o •I s a

^ to >> >• -c ^ i> E §&"> fe-» S ns O 1H >- a .=: •5 i

5 > . > "S.S5 II wps ;>*ps OK :/3pS :/)PS C/2PS P-.W U. tetralasmus X X X X X X X X C. pa rv a X X X X X X X L. claibornensis X X X X X X V. vibcx X X X X X X X V. licnosa X X X X X X X X L. anodontoides X X X X X X X A. perplicata X X L. excavatus X M. hembeli X A. suborbicaJata X

Nine species of the Alabama-Coosa River Rafinesquc (figured by Ortmann, 1919: to have 2 the Interior Basin system (Table 1, B) appear 95, pi. 8, fig. ) reached reached the Apalachicola River through a from the Alabama-Coosa River system. If former stream confluence (Text-fig. 1, see this can be shown, then EUiptio s. s. is a p. 285), since they are mostly missing from genus originally from the coastal regions. intervening systems. With the probable Four species of Unionidae appear to exception of Anodonta grandis, A. imbe- have reached the Apalachicolan region cilis, and Glebula rotundata, it is possible from a former confluence of the Apalach- that the remaining six species were once icola and Savannah river systems (Table endemic to the Alabama-Coosa River 1, C). They will be discussed with the system. Matteson (1948a: 131) suggests Southern Atlantic Slope fauna. Three ad- that EUiptio c. crassidens and E. dilatata ditional species entered the Savannah

Table 1, B. Unionidae, derived from the west, found in both the Alabama- Coosa and Apalachicola river systems, supporting the assumption of a former confluence between them.

p n> a U u a v a *-* .3- .a . a >> M a "> ^ a O u a •a5 a in o §. S.fe J3.2 WPS >HpS UPS Ops c/3PS E. crassidens X X X A. boijkiniana X X X A. grandis X X X A. imbecilis X X X E. arctata X X X X A. radiatus X X X S. subvextis X X G. rotundata X X X M. penicillatus X X 272 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Table 1, C. Unionidae, derived from the east or west, found in the Alabama- COOSA AND APALACHICOLA RIVER SYSTEMS, AND WHICH GIVE EVIDENCE OF A FORMER CONFLUENCE WITH THE SAVANNAH RlVER SYSTEM, EXCLUDING THE POSTULATION OF TWO MINOR STREAM CONFLUENCES. Atlantic Slope Uniomdae • Johnson 273

Table 1, E. Unionidae endemic to individual riveb systems IX the Apalachicolan region. 274 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Table 2. Distribution' of the Unionidae of the Atlantic Slope region from the altamaha to the potomac river systems, and relevant species FOUNT) IN THE ApALACHICOLA RlVER SYSTEM.

> C 5 -c £ C £ § £ 1* a >

s a < Atlantic Slope Unionidae • Johnson 275

Table 2, A. Unionidae, most of which are widely dim huh n i> in the Northern Atlantic Slope region, all but one of which terminate in the Southern Atlantic Slope region, but are of ddverse origins.

£ £ £ s > s CD 0) p. u Vj «n -5 >> >. 6 >. ©I. I- en a w fct, XB O en X J- S * O I. — - •n d e S* i- O CD s© 7 > >2 5 > Ok -- >': « Ck

A. implicate L. nasuta A. heterodon X L. radiate radiata X A. undulata X X S. undulatus X X L. suhviridis X X X A. varicosa X X L. cariosa X L. ochracea X X X 276 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Table 2, C. Unionidae derived from the Apalachicolan region, found in the Savannah River system, which afford evidence of a former confluence with the apalachicola rlver system, and which terminate in the Southern Atlantic Slope region.

4)

S«i £s o» (3 y V II 2 ' JB R, (D £ * O j- Zj & O OJ X > "3 > -1 > 2*> > h El le- u£ Sutf ££ C;£ s:£ ££ Atlantic Slope I'momdak • Johnson 271

Table 2, E. 278 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Table 2, F. Unionidae endemic to individual river systems of the southern Atlantic Slope region.

a 4) a> o> ^

Pj ^ s > c £ 'O > S-S 8.1&£ g-S -5-S <& Otfi c/s« Wrt IS

£. waccamawensis Atlantic Slope Uniomdak • Johnson 279

Table 3. The Unionacea of the Northern Taule 3, A. I'mionidae eoi \n i\ the Noiitukkn Atlantic Slope region. Atlantic Slope region with close affinities to species of the interior b\m\, believed to have come around the northern end of mil 1. Margaritifera margaritifera (Linnaeus) Appalachian Mountains. 2. Ettiptio complanata (Lightfoot) 3. Ettiptio lanceolata (Lea) Basin 4. Lasmigona subviridis (Conrad) Interior Northern Atlantic Slope 5. Alasmidonta undulata (Say) E. dilatata E. complanata 6. Alasmidonta marginata (Say) A. marginata A. varicosa 7. Alasmidonta varicosa (Lamarck) A. grandis A. cataracta cataracta 8. Alasmidonta heterodon (Lea) A. grandis A. cataracta fragilis 9. Anodonta cataracta cataracta Say A. grandis A. implicata 10. Anodonta cataracta fragilis Lamarck L. ovata L. cariosa 11. Anodonta implicataSay L. ovata L. ochracea 12. undulotus Strophitus (Say) L. radiata sUiquoidea L. radiata radiata 13. Ligumia nasuta (Say) 14. LampsUis cariosa (Say) 15. LampsUis ochracea (Say) 16. LampsUis radiata radiata (Gmelin) cated that the predecessor of complanata reached the Savannah River system from the Alabama-Coosa River system by stream not spread significantly into the region, are capture. In Chapter 1 of this paper, a dif- not discussed. ferent order of stream capture is postu- Ortmann (1906, and 1913a: 364) sug- lated. It appears that the Apalachicola and gested that the ancestors of the Atlantic Savannah river systems were connected Slope Unionidae listed above, with the ex- and separated before the Apalachicola and ception of Elliptio complanata (Table 3, Alabama-Coosa were joined, judging from A), migrated around the northern end of the distributional evidence afforded by the the Appalachian Mountains, in preglacial unionid fauna. The present distribution of which times, by way of the Eirigan River, complanata in the Apalachicolan region, flowed in the direction of the St. present as it is restricted to the Apalachicola River Lawrence River. There was no barrier to system, also indicates this order of stream their southward because the base- dispersal capture. It was probably already a species leveled coastal then extended much plain before its ancestor, unidentified by Matte- farther seaward. son, was supposed to have arrived on the During glacial episodes of the Pleisto- Atlantic Slope. E. complanata is related to cene, the species found on the eastern side dilatata (figured by Ortmann, 1919: 95, pi. of the mountains were effectively separated 8, fig. 2) of the Interior Basin. Both spe- from those of the Interior Basin, and speci- cies enjoy a certain ubiquitousness within ation took place among them, probably their areas of distribution. The history of south of the limit of glacial drift, early in similar complanata is, in short, probably the period. Later some of these species to that of the other species discussed spread into the Southern Atlantic Slope above. 1 region where the coastal region was base- leveled. After the Pleistocene, this assem- 1 Walker (1910: 131) suggested the same origin blage of species occupied most of the of E. complanata as that presented here. He northern lost its territory by predecessors. further thought that L. sabrostrata and L. nasuta, Ortmann (1913a: 361) was unaware of and A. calceolus (figured by Clarke and Berg, and A. heterodon had similar the complete distribution of E. complanata 1959: 29, fig. 30) relationships and the same origins as the species and suggested that it had its center of pairs discussed above. However, Ortmann's argu- Southern At- radiation somewhere on the ments as to their origins are thought to be more lantic Slope. Matteson (1948a: 131) indi- trenchant and are followed here. 280 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Table B. Unionidae found in the Atlantic Ortmann (1919: 109) called attention to 3, Slope region which appear to have crossed the the presence of complanata in the small divide of the Allegheny Mountains, or are of the Sus- tributaries of the West Branch represented by allopatric species. quehanna River, Pennsylvania, close to the divide, and to dilatata in the Allegheny Interior Basin River drainage, but he did not suggest that dilatata entered the Atlantic Slope here and speciated. He did, however, suggest that Strophitus undulatus and Alasmidonta marginata entered the Atlantic Slope here. These species are discussed later. The Margaritiferidae is represented on the Northern Atlantic Slope region by a single species, Margaritifera margaritifera. It is a special case because of its origin, but it is included here since its post-glacial history is similar to that of the other species discussed above. There is good evidence that M. mar- garitifera (figured by Ortmann, 1919: 2, 1 the Pacific pi. 1, fig. ) reached region from Asia during the Miocene or early Pliocene when the two continents were connected by a North Pacific (Bering) bridge (Walker, 1910). Walker further speculated that this species separately reached the Atlantic Slope by an Atlantic bridge over Iceland and Greenland, prob- 1 ably at roughly the same time. In any event, M. margaritifera appears to have reached eastern North America before the Pleistocene, during which time its range was restricted. Ortmann (1913a: 378) found it in Pennsylvania just south of the limit of glacial drift and suggested that Pennsylvania was at least one re- fugium where M. margaritifera survived the Pleistocene Epoch. Its present distri- bution, from Pennsylvania to Newfound- land and Labrador, is the result of a Atlantic Slope Unionidae • Johnson 2NI

verv close to Villosa vanuxemensis (Lea), bers of the genus found in the Interior resemble one another. (for reference to a figure see Simpson 1914, Basin, nor do they 1: 165) which is found in the Holston These two Alasmidonta appear to have River and elsewhere in the Tennessee River long existed on the Southern Atlantic Slope, but distributed above the system. It is not obvious where the an- are now widely cestor of constricta came from; it may have drift border. entered the Roanoke from the New River 4 along with L. suhviridis, if it can be as- CHAPTER was more sumed that vanuxemensis widely The generic affinities of the Unionacea time. distributed in pre-glacial of the Apalachicolan region, Peninsular Alasmidonta Strophitus undulabus and Florida, and Southern and Northern At- crossed the divide in central marginata lantic Slope regions. Pennsylvania, in the vicinity of Cambria, Westmoreland, and Indiana counties, Summary where the Susquehanna drainage of the Over forty genera of Unionidae occur in east has encroached on the Alle- largely the Interior Basin, many of which are also River of the west, and where stream gheny found in the Alabama-Coosa River system. has taken place (Ortmann, 1913a: capture In the Apalachicolan region are fourteen A. remained restricted to 368). marginata genera that occur in the Interior Basin as the River while upper Susquehanna system, well as in the Alabama-Coosa River system. S. undulatus throughout the Atlantic spread An additional genus, Glebula, is limited to region. Both of these species go into Slope the Coastal regions. In the Apalachicolan small streams and, in general, avoid very region also occur Margaritifera, Alasmi- large rivers. latter is donta s. s., and Quinctincina. The nasuta is with L. Ligumia allopatric endemic. The six genera found in Penin- subrostrata reference to a Say (for figure sular Florida occur in both the Apalach- see 1914, 1: 99) of the central Simpson, icolan and Atlantic Slope regions but are and western part of the Interior Basin. It probably derived mostly from the former. probably developed in the lake drainage Nine of the eleven genera of Unionidae of Ohio in postglacial time and spread found in the Southern Atlantic Slope region eastward into the Hudson River, following are found in the Apalachicolan region. The a course along the baseleveled coastal exceptions are Lasmigona and Ligumia. plain. It reached the James River system Four of the to be of south- in the south and the St. Lawrence River genera appear affin- ern origin, and the remainder have system, Canada, in the north (Ortmann ities with both the north and south. Only 1913a: 379). This species has a preference seven of these genera occur in the North for quiet water. Atlantic Slope region. This region is also Elliptic) lanceolata occurs in the Apa- which does not lachicolan region and on the Atlantic Slope occupied by Margaritifera, occur on the Southern Atlantic Slope. but there it extends only to the Susque- Ortmann 323), speaking of the hanna River system, Pennsylvania. The (1913a: the Atlantic is a Unionidae that reached Slope present distribution argues that this an- around the northern end of the Appala- southern species, but its immediate noticed the diminution cestors or the area of its speciation is chian Mountains, and con- obscure. of genera from the Interior Basin, Mountains Alasmidonta undulata and Alasmidonta cluded that "the Allegheny formed an barrier to the east- heterodon appear to be relicts of a much important ward distribution of the bulk of the west- older, westerly derived fauna, since they is noted here that the no longer closelv resemble anv of the mem- ern fauna." It along 282 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Table 4, A. The genera of Unionacea in the Table 4, C. The genera and subgenera of Apalachicolan region. UNIONroAE IN THE SOUTHEBN Atlantic Slope region.

Margaritifera? Anodontoides Fusconaia Strophitus Pleurobema s.s. [Not represented in this area] 4 Ambelma2 Obovaria Lexingtonia. Endemic to the Southern Atlantic Quincuncina Carunculina Slope Pleurobema Villosa Elliptio s.s. to Altamaha River Ettiptio Lampsilis Canthyria. Endemic the sys- Uniomerus Medionidus tem Alasmidonta3 Glebula Uniomerus Anodonta Lasmigona s.s [Not represented in this area] Platynaias 1 See note on p. 280. Alasmidonta s.s. - See note on p. 270. 3 Alasmidonta is represented in the Alabama-Coosa Prolasmidonta. Endemic to the entire Atlantic River system by A. mccordi Atheam (1964: 134, pi. 9, figs, a, b). Slope 4 Obovaria is found in the Alabama-Coosa River system Decurambis and is represented in the Apalachicolan region only in the Anodonta s.s. in this area] Escambia River, the next system immediately east of the [Not represented Alabama-Coosa River system. Pyganodon Utterbackia Strophitus southern coastal plain, where no such great Carunculina barrier existed, there is a similar diminution Villosa of genera from west to east. Ligumia Lampsilis s.s. It can be seen in Table 1 that fifteen of the seventeen genera found in the Apa- lachicolan region (Table 4, A) occur in included here in the Apalachicolan region.) the Alabama-Coosa River system. The In this area occur only six genera (Table exceptions are Margaritifera (represented 4, B ) derived from the west and north-east. by M. hembeli, a relict), which is also found Of the eleven genera ( Table 4, C ) found in one other to the west, drainage in the Southern Atlantic Slope region, nine the and Bayou Teche, Louisiana, Quincun- are represented in the Apalachicolan region is the cina, which endemic to Apalachicolan where there is a total of seventeen genera. Alasmidonta occurs in region. Although Strophitus, Lasmigona, and Ligumia are it does the Alabama-Coosa River system, clearly from the North. Of these, Stro- not appear close to Alasmidonta s. s., which phitus alone is represented in the Apalach- is found only in the Apalachicolan and icolan region by S. subvexus (Conrad), Atlantic Slope regions. which is endemic to the region. Peninsular Florida is considered to be Of clearly southern origin are Uniomerus the area south and east of the Suwannee and Carunculina. Pleurobema and Villosa River system and south of the St. Marys are probably of southern origin. The genera River system. (Both of these systems are Elliptio, Alasmidonta, Anodonta, and Lampsilis have affinities with both the southern and northern elements of the Table 4, B. The genera and subgenera of Umonidae in Peninsular Florida. fauna. eleven Table found Of the genera ( 4, D ) Elliptio s.s. in the Southern Atlantic Slope region, seven Uniomerus occur in the Northern Atlantic Slope region, Anodonta s.s in this area] [Not represented where there also occurs an additional Utterbackia genus Carunculina of Unionacea, Margaritifera. These eight Villosa genera repopulated the entire Northern s.s Lampsilis Atlantic Slope above the limit of glacial Atlantic Slope Unionidae • Johnson 283

and Table 4, D. The genera subgenera of mountains, running first between the Unionacea in the Northern parallel ridges in the Atlantic Slope region. (anticlines) synclinal valleys, and finding their outlets in a north- westerly direction toward the Interior Margaritifera Basin. On the Atlantic side of the Elliptio s.s divide, Lasmigona s.s shorter streams originated in the highest Platynaias elevations and flowed east and southeasl Alasmidonta s.s. across the Archaic rocks. Rivers also Prolasmidonta. Endemic to the Atlantic Slope formed on the tops of the anticlines, and region Decurambis anticlinal valleys developed, parallel to the Anodonta s.s. [Not represented in this area] synclinal ones. The uppermost rocks of the Pyganodon anticlines were largely hard sandstones of Strophitus Carboniferous age, below which were Ligumia softer shales and limestones of Devonian Lampsilis s.s. age or older. The synclinal rivers ran uni- formly over the same hard sandstone, but the anticlinal those on the drift, with the exception of Lasmigona, rivers, primarily were the first to cut which only reaches the Hudson River highest elevations, the the rivers system, New York. One of these, Ligumia, through sandstone, allowing access to the softer rocks below. In appears to have entered from the west at time, the close of the Pleistocene. With this the anticlinal valleys became more exca- vated than the This exception, the other genera were estab- synclinal valleys. advanced farthest in the eastern lished below the limit of glacial drift be- process section of the mountains, so that what was fore that period. once the highest elevation became a deep CHAPTER 5 valley. By the Cretaceous Period, nearly all of The relationship of the geomorphology the Appalachian area was peneplaned, with and topography of the Apalachicolan and the exception of a chain of monadnocks Atlantic Slope regions to their unionid between eastern Tennessee and North faunas. Carolina, and scattered hills in northern New England. (These unreduced areas The Mountains have been Appalachian form the modern Great Smoky Mountains barrier the of Union- an effective to spread and the summits of the White Mountains in acea from the Interior Basin to the coastal . Elsewhere there are area since time postcretaceous (Ortmann, widespread remnants of the peneplane 1913a: 383), although the original Appala- even on the crests of the highest present chian were formed much earlier Mountains ridges.) The coastal plain was covered in the Permian Period as the result of with a shallow sea that deposited a veneer lateral pressure. They consisted of a num- of beds on the Archaic rocks. Because of of ber of parallel anticlines and synclines that certain characteristics of the patterns ran in a northeast-southwestern direction. some of the present streams, it is thought inland than These folds were pressed against an old that this sea extended further block of Archaic rocks, the present Pied- is now indicated by existing beds. of the mont Plateau, that lay to the east of them. The relief present Appalachian is almost to a series The highest elevation of these folds was Mountains due wholly that occurred in the south, and the divide was well to of broad regional upwarps the the east, close to the old Archaic land. in the early Cenozoic Era, giving pres- The rivers followed the structure of the ent elevation both to the already extant 284 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

mountain structures and to the subsequent and clay. Waterfalls are so numerous at sculpture of this complex mass by erosion. the boundary between the Piedmont The Appalachian Mountain system of east- Plateau and the Coastal Plain that it is ern North America extends in a wide belt, known as the Fall Line. During the Pleisto- more or less parallel to the coast line for cene, a number of changes in the level of more than 1600 miles from the Gulf Coastal the ocean left terraces on the Coastal Plain in central Alabama to the St. Law- Plain that will be discussed below. rence River, Quebec. The mountains are The whole Appalachian Mountain system divided from west to east into four prov- has also been divided from south to north 1 or into three Southern. North- inces. ) The Cumberland, Appala- regions. 1) chian Plateau, which extends from Ala- eastern Georgia to the New or Kanawha bama to New York and merges with the River, Virginia, and . 2) Allegheny Plateau. This western area has Central. New River to the Hudson River been peneplaned, but the soft rocks of valley (covered here to the Potomac River pre-Carboniferous age have not been system). 3) Northern. Hudson River val- reached. 2) The Appalachian Valley and ley to the lower St. Lawrence River. This Ridges that reach from Alabama to New adumbration of the complex history of the York. In the south, this province has a Appalachian Mountains, primarily of the valley-like appearance, but in the north, it area under study, is offered as background. becomes the Allegheny Mountains. The The relevant details follow. eastern part of these were originally much The main backbone of the present Appa- higher. They are cut into by anticlinal lachian Mountains extends continuously streams, which have carved out broad from the divide between the Toccoa and limestone valleys, with high ridges of Coosawattee rivers in northeastern Georgia harder rock between them. Within the to northern Virginia, and is known as the Allegheny Mountains, and parallel to them Blue Ridge. From it the streams flow on the east, where there was once the northwest, southeast, or east. It has been highest elevation, there is now the ex- mentioned that the original divide was well ceptionally broad Great Allegheny Valley. to the east, as is the present divide, and 3) The Blue Ridge is the highest part of that the eastern streams did, and do, have the Southern Appalachian Mountains and a steeper grade and a shorter, more direct extends from northeastern Georgia across route to the sea. Since they had, and have, western South and North Carolina into more eroding power than the streams of southwestern Virginia. The Blue Ridge of the western drainage, they have encroached the Central Appalachians extends from on it. There has also been a tendency of Central Virginia into Pennsylvania where the rivers on the Atlantic Slope to capture it continues as South Mountain. It is the streams between systems. flank of an anticline, consisting largely of Evidence of stream capture. The southern Archaic rocks. It is less elevated here than extremity of the Blue Ridge is the water- in the south. 4) The Piedmont Plateau shed of a number of rivers. The Toccoa, (or Upland) goes from central Alabama the Little Tennessee, and the French Broad to the Hudson River, New York. all flow into the Interior Basin. Of concern To the east of these mountain provinces here are the two systems that originate on lies the Coastal Plain; this is a continuation the southeast side of the mountains and of the Archaic granites of the Piedmont that flow into the Gulf of Mexico; the Plateau, over which are much younger Etowah River of the Alabama-Coosa River Cretaceous and Tertiary deposits of sea- system, which flows westward around Pine lain limestone and poorly consolidated , Georgia, into the Appalachian Atlantic Slope Unioniuai. • Johnson 285

Valley and then into the Gulf; the Chatta- Paris Mountain, then flows eastward and hoochee River of the Apalachicola River northeastward. It is clear that the Roanoke system, which flows sonthwestward into River has captured here a tributary of the the Gulf; and the Tugaloo River of the New River. The presence of La.smigona Savannah River system, which also heads subviridis and Villosa constricta in the in the mountains and flows southeast into Atlantic Slope region adds credence to the the Atlantic Ocean. On the Piedmont geomorphological evidence. Plateau, as can be seen in Text-figure 1, Beginning with the Roanoke, the suc- (see p. 268) all three of these rivers have ceeding river systems, the James, Potomac, tributaries that presently are separated by and Susquehanna, all flow more directly minor divides. Matteson (1948a: 131) sug- east and each one cuts farther back into gested that confluence occurred among the mountains (Campbell, 1896: 675). these streams during the late Tertiary. It Campbell also assumes that stream capture may have been later, but in any event, the has taken place between the headwaters distribution of the Unionidae in the several of the Roanoke and James river systems, river systems indicates that confluence did an assumption which might explain the take place. presence of Villosa constricta in both A number of other Atlantic Slope river systems. systems, in addition to the Savannah, have The Potomac River cuts across the their sources near or in the Blue Ridge. mountains and has, in northeastern West The Cooper-Santee River system, which Virginia, reached the Allegheny Plateau. drains some 15,000 square miles of central It now drains a longitudinal synclinal val- South Carolina, has three main tributaries, ley. Campbell (1896) further states that the Saluda, Broad, and Catawba, all of in the mountains, the Potomac River which originate close to the divide. The robbed the James River of tributaries, and Pedee River system's main tributary, the that the Susquehanna River of Pennsyl- Yadkin, begins in Watauga and Caldwell vania encroached on the Potomac. How- counties, North Carolina, near the divide. ever, these sequences of stream captures The Roanoke River system, which has the seem too ancient to be relevant to the Dan River as a major tributary, forms in present unionid fauna north of the James. Patrick County, Virginia. The Roanoke it- Many of the Unionidae appear to have self commences in the eastern foothills of spread along the baseleveled coastal plain; the Blue Ridge. this spreading is discussed below. It is only in the southern Appalachians The Piedmont Plateau. In Alabama, the that streams that flow westward still have Piedmont Plateau is divided into two their sources near, or in, the Blue Ridge. regions. 1. The Ashland Plateau, the high- The best example is the New River, which est division, is an area of crystalline rocks, preserves its ancient course and cuts across diversified by ridges. It is the mountainous the divide. According to Campbell (1896: portion of the Piedmont and includes the 674) there is a good example of stream , the highest point in of the capture in the uppermost Roanoke River, state. It is drained by portions River 2. The Virginia (Plate 1). Alabama-Coosa system. Ope- The headwaters of the Roanoke's North lika Plateau is lower and is underlain by Fork run first in a southwesterly direction Archean rocks; it has no striking topo- features. It is drained the in a valley that clearly continued to New graphic by tributaries of the River of the River; but just north of Christiansburg, Tallapoosa which lies Montgomery County, Virginia, the North Alabama-Coosa system, adjacent border of the Archaic Fork makes a sharp bend, cuts through to the northwest 286 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

rocks, and by the Chattahoochee River, complanata, E. congaraea, and Villosa which forms the eastern boundary of the delumbis, which are found in the Broad state. On the Opelika Plateau the rivers River, opposite to which, in Abbeville have cut valleys some 200 feet below the County, South Carolina, have also been general surface. The region continues into found Pleurobema masoni, Elliptio fra- Georgia as the Greenville Plateau, and is tema, E. arctata, Lasmigona subviridis, and drained by longitudinal streams that flow Lampsilis cariosa. southwestward and westward to the Gulf. The siliceous character of the Piedmont The remaining portion of the Piedmont of water, as illustrated in Text-figure 4, might Georgia to the north is drained by dendritic offer an adequate explanation for the dim- streams. Those rivers that originate in the inution of the fauna above the Fall Line. Piedmont of Georgia include: the Flint of However, elsewhere on the Atlantic Slope the Apalachicola system, which flows into siliceous water does not appear to be such the Gulf, and the Ocmulgee and Oconnee an inhibiting factor to the distribution of of the Altamaha system and the Ogeechee the Piedmont Unionidae. system, which both flow into the Atlantic. The Coastal Plain. The Coastal Plain is In South Carolina the rivers have their a continuation of the Archean rocks of the sources in the mountains or on the Coastal Piedmont over which occur Cretaceous and Plain. In North Carolina the Piedmont Tertiary deposits of sea-lain limestone and rivers include: the Cape Fear, Neuse, poorly consolidated sand and clay (Text-

in Vir- . Pamlico, and Chowan systems; and fig. 2 ) In spite of the apparent propinquity ginia, the York and Rappahannock systems. of the Apalachicola and Altamaha river Ortmann (1913a: 340) observed, in the systems in the Piedmont where the Chatta- area he studied, that the great majority hoochee passes close by the source of the of Atlantic Slope Unionidae go up into the Ocmulgee, the unionid fauna does not mountains (where the rivers may have indicate any direct exchange of species. reached Cretaceous limestone) and ap- The Flint River has no large tributaries proach the headwaters without significant to the east, and in some places it is as depauperation. much as fifteen miles from the divide that In the Apalachicola River system, pres- separates it from the drainage to the east, ent pollution prevents verification, but though Big Indian Creek, near Fort Valley, older records indicate that some of the Houston County, a tributary of the Ocmul- unionid fauna of the Chattahoochee River gee River, is only three miles distant. It extended on the Piedmont at least to the would appear that the height of the Tifton vicinity of Atlanta, Georgia. In the Flint Upland and Fort Valley Plateau, and the River, which forms south of Atlanta, depth of the river channel on the Dough- Unionidae are found in Line Creek, close erty Plain, have prevented stream capture to the headwaters. between these river systems for a long time. On the Atlantic Slope of Georgia the During the Pleistocene the Coastal Plain unionid faunas of the major river systems was subjected to a number of invasions by —the Altamaha, Ogeechee, and Savannah the sea. According to Cooke (1945: 248), —end abruptly at the Fall Line. The ex- the maximum and earliest flooding left a ceptions in the Altamaha River system are record, the Brandywine Terrace [Hazlehurst EAliptio complanata and Anodonta c. cata- in Georgia], which correlates with the rocta, which have been found near Stone Aftonian interglacial stage. The sea was Mountain, DeKalb County, the headwaters thought to be 270 feet above the present of the Ocmulgee River. In the Savannah level. As mentioned on p. 268, MacNeil River system the exceptions are Elliptio cast doubt on the extent of Afonian flood- Atlantic Slope Unionidae • Johnson 287

2. Text-figure The topographic divisions of the Coastal Plain of Georgia. (From La Forge, 1925, fig. 2.)

ing, and concluded that there is no evidence In spite of these valid objections, some of to indicate that the sea ever reached a level the terrace names are well established in more than 150 feet above the present one the literature and are retained here, but during the Yarmouth interglacial stage. He are used in the broader sense implied by regarded the Brandywine Terrace to be Alt and Brooks. of subaerial origin. Russell (1957: 427-428) On the basis of the geological evidence, suggested that the shore lines might have Alt and Brooks (1965) concluded that the been subjected to differential uplifting. 215-250 [270] foot shore line was occupied Oaks and Coch 1963 on the basis of the Miocene. This ( ) , during Upper dating cores made in Virginia, postulated six cy- was confirmed by Laesse (1968) with bo- cles of the Pleistocene seas, with maximum tanical evidence. Alt and Brooks also con- heights of 45 feet above present levels. cluded that the 90-100 foot shore line was with the Pliocene. Alt 1968 They disagreed the theory of "terrace- occupied during ( ) stratigraphy" and disapproved of the no- further concluded that the 70-80 foot shore menclature of the terraces, or shore lines. line also dated from the Pliocene. He rec- Alt and Brooks (1965) also objected to ognized three Pleistocene shore lines: one the nomenclature of the shore line scarps at 40-45 feet, which he assigned to the Af- and their associated terraces, since the im- tonian interglacial; a second at 2.5-30 feet, plication is that they refer to independent which he assigned to the longer Yarmouth physiographic units rather than to com- interglacial; and a third at 5-10 feet, which ponent parts of a single physiographic unit. he assigned to the Sangamon interglacial. 288 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

oVienna. tericds

\ «Cordelp ''Georgetown if «Cuthberto[)aw30n S *Lec»burp °Ashburn

[ATbnx.y OcFlla. ••Sylvester Tifton A;

^'•C 1 1 a. ami <—J-)°No_5hvifle

Wiwfe—IE—it—Jfe—JTw 'lib

A. Text-figure 3. The Coastal Terraces on the Coastal Plain of Georgia. (From Cooke, 1925, pi. 10, fig.

The coastal terraces of Georgia are gion, since all of the larger rivers have ter- shown in detail in Text-figure 3 because races of their own. A river terrace may be some are especially relevant. These ter- defined as the old floor of a valley within races were formerly defined as sea bottom which an inner valley has been cut by a uncovered and converted into land by a stream when deepening its channel. Thus, drop in sea level, but Hoyt and Hails river terraces are fossil flood plains. 1967 these terraces The various coastal terraces shore ( ) now regard Georgia (or as former lagoon salt marshes. The terraces lines) are correlated below from oldest are not entirely confined to the coastal re- (top) to youngest:

MacNeil Alt and Brooks (1965) Murray (1961: 508) Cooke (1925: 35) (1950: 99) and Alt (1968)

General maximum height Georgia Range Florida Time or Stage

Brandywine 272 feet Hazlehurst 270-215 feet Upper Miocene 250-215 feet Coharie 215 Claxton 215-160) Okefenokee Pliocene 100- 90 Sunderland 170 Okefenokee 160-100J

Wicomico 100 Penholoway 100-60 1 IVnholoway 70 Wicomico Late Pliocene 80- 70 42 Satilla 60-0 Aftonian 50- 45 Talbot J interglacial Pamlico 25 Pamlico Yarmouth interglacial 30- 25 Sangamon interglacial 10- 5 Atlantic Slope Unionidal • Johnson 289

4. Text-figure Water types of Georgia. (From Cherry, 1961, fig. 6.)

A number of the Coastal Plain rivers Among the river systems that are com- appear to have their sources above the pletely on that part of the coastal plain that level of maximum flooding by the sea; was flooded during the Pliocene is the among these are the Escambia, Yellow, St. Marys. Text-figure 3 shows that it is Choctawhatchee, Ochlockonee and Su- entirely on the Olcefenokee and Penholo- wanee river systems in the Apalachicolan way (Wicomico) formations. The river region and the Satilla and Edisto river sys- drains the Okefenokee Swamp, and its tems on the Atlantic Slope. The Yellow, water, with a low pH, is high in chlorides Satilla, and Edisto river systems appear to or silicates (Text-figs. 4 and 5). Although have been more effectively inundated by the Satilla River is similar, it differs from the sea, because their small unionid faunas the St. Marys in that a small portion of its suggest subsequent repopulation. headwaters is on the Tifton Upland, be- 290 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Text-figure 5. Ranges in pH of Georgia water. (From Cherry, 1961, fig. 8. yond the area of maximum Pliocene and as the St. Marys. It is completely on Pleistocene flooding. Both rivers have a the Wicomico and Chowan formations and small unionid fauna (Table 1). The domi- has a typical coastal assemblage of Unioni- nant species in the St. Marys is E. c. crassi- dae (Table 2), but it is remarkable in dens, and in the Satilla, it is E. crassidem having an endemic species, E. uaccomo- downiei. If the Satilla was a refugium, it ivensis. is possible that downiei had a chance to Among the small river systems of more evolve there and that E. c. cra.ssidens came recent origin on the Coastal Plain in the back from the west by a route not explained Apalachicolan region in which Unionidae here. have been found are the Econfina and the The Waecamaw River system in South Wasassasa. In the Econfina occur Elliptic and North Carolina is of the same origin icterina, ViUosa lienosa, V. vibex, and Atlantic Slope Unionidae • Johnson 291

Lampsilis anodontoides. E. icterina occurs CHAPTER 6 in the Wasassasa. and Conclusions The systems of more recent origin on the Summary Coastal Plain in the Atlantic Slope region The Apalachicolan region as defined are the Combahee, Ashley, Black, and herein includes not only those river systems rivers. White Oak The Combahee has five flowing into the Gulf of Mexico, from I lie species: E. complanata, E. icterina, Unio- Escambia to the Suwannee, but also the merus tetralasmus, V. delumbis, and V. St. Marys and Satilla, which flow into the vibex. In the other river systems occur Atlantic Ocean. Forty-nino species com- either, or both, E. complanata and icterina, prise the unionid fauna of this region. with the exception of the Black, in which Nineteen have affinities with species to U. tetralasmus also occurs. the west or Interior Basin, twelve are en- It is generally regarded that the Pleisto- demic to the region, and eleven are re- cene lasted for a million years, although stricted to individual river systems. Three Flint (1957: 301) states that, "all we can species that were probably originally en- say with confidence is that the last major demic to the region have spread into the glaciation occurred within 30,000 years, Atlantic Slope region, and four others and on the basis of sea floor stratigraphy, probably originated on the Atlantic slope. the Pleistocene as a whole, embraces at The Atlantic Slope region is defined as least 300,000 years, perhaps much more." those streams flowing into the Atlantic In any event, during the Pleistocene at least Ocean from the Altamaha Biver system, five ice sheets pushed southward into the Georgia, to the lower St. Lawrence Biver United States from centers of accumulation system, Canada, including rivers in New- in Canada. The interglacial times are foundland and Labrador. Since the area is thought to have been of long duration with occupied by an assemblage of species of sea levels high. During the shorter glacial both southern and northern origin, it can times the sea must have been relatively be divided into Southern and Northern low, and the coastal plain often extended regions. The former extends north to the much farther to the east than it does at James Biver system, Virginia, the northern- present. most system with a clearly southern assem- Emery (1967) has convincingly shown blage of species. Forty species comprise that as early as 19,000 years ago (lowest the unionid fauna of the Atlantic Slope level of the sea during the Wisconsin region, but one, Lampsilis ovata (Say), was glacial stage) that this was so then. artificially introduced from the west at the Ortmann (1913a: 383) pointed out that turn of the century. Of the remaining there is a dispersal line directed both north thirty-nine species, sixteen occur in the and south on the Coastal Plain, because Northern Atlantic Slope region. Of the geophysical barriers have been largely re- twenty-three additional species found on moved by baseleveling. Lowland flooding the Southern Atlantic Slope, seven appear allows fishes to migrate from one system to to have been derived from the west, seven another. The present paucity of Unionidae are endemic to the region, and seven are in most of the coastal streams of recent endemic to individual river systems. Two origin indicates that dispersal was probably other species, thought to have originated easier during one of the inter-glacial on the Southern Atlantic Slope, are found periods. in Peninsular Florida, and one of them has The importance of baseleveling and its spread into the Apalachicolan region. faunal significance was well illustrated by 1. Because of their unique mode of dis- Adams (1901). tribution, the Unionidae are able to furnish 292 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

important evidence for past conditions of distinct, consisting of species clearly de- drainage by their present distribution, as rived from the fauna of the Interior Basin, was demonstrated by Ortmann (1913a: and a secondary, or Atlantic fauna (Simp- 381). The discontinuous distribution of the son, 1900: 505), originally of Interior Basin Unionidae of the Apalachicolan region origin, but whose ancestry is more remote. indicates that there was once a confluence 6. In the Atlantic Slope region, most of of the headwaters of the Alabama-Coosa the species that are clearly of Interior Basin and Apalachicola river systems. The pres- origin are a northern group that migrated ence of the same species and analogs in the around the northern end of the Appa- Savannah River system on the Atlantic lachian Mountains before the Pleistocene; Slope as in the Apalachicola River also there is also a southern group that entered indicates confluence. The absence of some the Atlantic Slope region through a con- of the species found in both the Alabama- fluence of the Apalachicola and Savannah Coosa and Apalachicola river systems indi- river systems, also in pre-Pleistocene time. cates that the confluence of the Apalach- 7. The large number of endemic unionids icola and Savannah river systems took in the Altamaha River system, Georgia, of place first. Two minor confluences are both old Atlantic and Northern Atlantic postulated: One between Uphauppee Slope origin suggests that this system is Creek of the Alabama-Coosa River system particularly old and that it was sufficiently and Uchee Creek of the Apalachicola River isolated during some of the Pleistocene to system, the other between the headwaters have facilitated speciation. of the Choctawhatchee River system and a 8. Dispersal in the Apalachicolan region tributary of the Chattahoochee River of appears to have been largely by stream the Apalachicola River system. capture. On the Atlantic Slope there is a 2. Contained in the fauna of the Apa- dispersal line, directed both north and lachicolan region are species that are found south, where barriers are rendered ineffec- in the Alabama-Coosa River tive 1901 . system, having by baseleveling ( Adams, ) The entered this system through a former con- rate of dispersal was undoubtedly influ- fluence with it and the Tennessee River enced by changes in sea level during the system. The region contains mainly en- Pleistocene. Although it cannot be specifi- demic species of Interior Basin origin. A cally demonstrated, stream confluence in few species have reached it from the At- the uplands also may have played a part lantic Slope Region. in the dispersal of the Atlantic Slope 3. The presence of endemic species, species. primarily in the larger Apalachicolan river 9. In the Appalachian Mountains, there systems, indicates that these species had are a few examples of species crossing the refugia above the maximum Pliocene and divide by stream capture. In addition to Pleistocene flooding, and originated before the confluence of the Apalachicola and Sa- then. vannah River systems, there were conflu- 4. Ortmann (1913a: 385) pointed out ences of the headwaters of the Roanoke and that the Allegheny Mountains formed an New rivers and of the Allegheny and Sus- old and well-marked boundary between quehanna river systems. Evidence of the aquatic of the Interior Basin and former existence of these confluences is those of the Atlantic Slope region since borne out by the present distribution of post-Cretaceous time. For the Unionidae some unionid species in the adjoining sys- this boundary includes the entire Appa- tems and by the physiography. lachian Mountain system. 10. Margaritifera margaritifera (Lin- 5. The Atlantic Slope unionid fauna is naeus), an ancient species from Europe, Atlantic Slope Unionidae • Johnson 293

had a refugium in Pennsylvania below from the St. Lawrence Basin to the Atlantic maximum Pleistocene glaciation, and Ligu- Slope in postglacial time. These are two mia nasuta (Say) appears to have migrated special cases of distribution.

PART II. A REVISION OF THE UNIONIDAE FROM THE ST. MARYS RIVER, FLORIDA, TO THE POTOMAC RIVER, MARYLAND.

INTRODUCTION Renaissance. This work was cited by Lin- naeus in 1758 for margariti- In of the title of this Margaritifera spite parochial 1 fera; by Lightfoot in 1786 for Elliptio section, I hold the belief, as do many evo- complanata; and by Gmelin in 1791 for lutionary biologists, that most faunal radiata radiata. It was not until studies are an anathema. Their authors Lampsilis January of 1817, when volume two of seldom accurately determine subspecies First American Edition the even Nicholson's of (Mayr, et al., 1953: 181) and often British Encyclopedia appeared, which in- their determination of species is suspect. cluded Thomas Say's article on Conchol- My excuse for this faunal approach is ogy, that many of the common Atlantic thought to be justified by the findings Slope Unionidae were described and presented in Part I. I have studied the figured. Eleven of Say's species are recog- Unionacea for almost superfamily thirty nized here. Many of these were redescribed years and each of the forty species covered two years later, in 1819, by Lamarck in vol- in this section has been mono- completely ume VI of his Histoire Naturelle des Ani- those whose ex- graphed, including range maux sans Vertebres. Only three of his tends the area studied. Although beyond names are recognized in this paper. In 1830 the of each is believed synonymy species and 1831 Isaac Lea of Philadelphia de- to be complete, and the modern species scribe several new species from the Southern concept has been assiduously applied, Atlantic Slope region, and in 1834 Timothy of is not claimed. If infallibility judgment A. Conrad published a little volume, New I am of creating composite species, guilty Fresh Water Shells of the United States, will be found among the they probably which included many of the species he had Nevertheless, if this ubiquitous Elliptic). found in the southeastern states. is the in Part I should not so, postulations A bitter battle arose between Lea and serious modification since most of require Conrad over the priority of some of the the species on which they are based are species, but their polemics, discussed by old and discrete to satisfy a sufficiently Wheeler (1935), are only of historical in- typologist. terest. Lea continued to describe Unioni- dae from the Southern Atlantic Slope until the col- History of the taxonomists and the 1874, by which time he had burdened lectors of the Atlantic Slope Unionacea. literature concerning the few species found The Taxonomists there with over one hundred and fifty- names. In this nine of Conrad's taxa Three species of Unionacea found in the paper, of Lea's are considered valid. Atlantic Slope region were figured in the and fifteen the the demise of Isaac Lea, the de- first printed book devoted entirely to With Unionidae of the south- study of mollusks. This was the great folio scribing of the was volume by Martin Lister, Synopsis Method- eastern states, exclusive of Florida, icae Conchyliorum, which appeared in 1 London in 1685, at the close of the English See p. 314. 294 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

essentially over. Between the years 1883- expedition to the Southern Atlantic Slope

1934, the Wrights, father and son, de- of Georgia was made by W. J. Clench, K. scribed fifty-two species of mollusks, most J. Boss, and S. L. H. Fuller in 1962, for the of which were from Florida (Johnson, Museum of Comparative Zoology. Since a of un- Ruth Patrick 1967 of the 1967a). The Wrights were pair then, ( ) Academy sophisticated naturalists who redescribed of Natural Sciences of Philadelphia has had many species, and aside from Villosa J. Bates and S. L. H. Fuller make collec- villosa, which is described in this paper, tions of Unionidae in the Savannah and their other relevant taxa are included in other Atlantic Slope river systems, in con- the various synonymies. nection with her limnological studies. In 1950-51 W. M. Walter made an ex- The Collectors tensive survey of the Neuse River system, North Carolina. This di- The first serious student of fresh-water collection was vided the Museum of mollusks to make a collecting trip to South among Comparative United States National Carolina and Georgia was Timothy A. Zoology, Museum, and of of Conrad. His route of 1833 is shown in de- Museum Zoology, University tail by Wheeler (1935: 26, 27). Before Michigan. and after this time, most collections were In the Carolinas and northward, collect- made by interested local residents who ing has been adequate, but not organized sent their material to Isaac Lea, and oc- in a way that can be discussed. Many stations have been made casual collec- casionally to Conrad, for description. In by the 1830's Lea received Unionidae from tors, and Ortmann collected systematically in Lewis Leconte, J. H. Couper, Professor Virginia. Shepard, and Edmund Ravenel. These men made it possible for Lea to THE PRIMARY SYSTEMATIC STUDIES OF describe most of the endemic species in the APALACHICOLAN AND ATLANTIC SLOPE

Altamaha River, Georgia. Later, Dr. J. P. UNIONACEA. Barratt sent him shells from the Abbeville Charles T. Simpson's Notes on the District of South Carolina. From the Caro- Unionidae of Florida and the Southeastern lina's Lea received, in his later shells years, States (1892) was the first attempt at a from E. Emmons and from C. M. especially revision of any of the species of the South- Wheatley. ern Atlantic Slope region. This work was No extensive took on the collecting place followed (1900) by his Synopsis of the Southern Atlantic Slope after Lea's time, Naiades, which was expanded into the until W. J. Clench and P. Okkelberg made Descriptive Catalogue of the Naiades a trip to Georgia in 1929. This trip was (1914). These two works include a re- followed by another by Clench and H. van vision of the Naiades on a worldwide basis. der Schalie in 1932; and then by Clench, Simpson's species concepts, in so far as C. V. MaCoy, and II. D. Russell in 1934. they concern the Apalachicolan and At- The collections made in 1929 and 1932 lantic Slope regions, have been generally were shared by the Museum of Compara- accepted, with but few changes, until tive Zoology and the University Museum, recently. His were scholarly works. Simp- University of Michigan. The remaining son was among the more competent mala- collections are in the Museum of Compara- cologists of his time. He included in his tive Zoology. In 1937 van der Schalie made works almost every taxon relevant to the a survey of the mollusks of the Ogeechee North American Unionacea that had ap- River, Georgia, for the Museum of Zoology, peared before 1914. University of Michigan. The last major After Simpson's work, aside from some Atlantic Slope Unionidae • Johnson 295

studies by Ortmann, little was done on the "Many generic divisions have Keen proposed, based characters of diverse some Unionaeea of the Apalachieolan region upon kinds, being those of the hard parts, others those of until van der Schalie wrote the Naiad the soft parts. These characters resemble Fauna the River, in North- of Chipola each other in the respect that they all vary western Florida (1940). This was followed very much, and that none are constant. The by Clench and Turner's Freshwater Mol- general plan adopted herein is to select groups which may he typed by one member, lusks of Alabama, Georgia, and Florida the rest being evident kinfolk. .Such new the Escambia to the Suwannee River from genera as are named herein are unaccom- (1956). Further additions were made by panied by diagnoses, a type shell being Athearn in Three new unionids from Ala- named, whose chief characters will forever such bama and Florida and a note on Lampsilis give diagnoses."' 1964 and in Additions jonesi ( ) by Johnson The paper is sprinkled with numerous to the unionid the drainage fauna of Gulf generic and subgeneric names. It is not and Florida of Alabama, Georgia (1967b). unfair to say that most of these names The Unionaeea of the Northern Atlantic would have fallen into obscurity except were Slope region elegantly monographed for two events. In 1934, Thiele, in his who wrote the Naiades by Ortmann, of Handbuch der Si/stematischen Weichtier- 3 (1919). Recently, Pennsylvania, pt. kunde (2, pt. 3: 815-844), fell under both Clarke and made a modern Berg (1959) the systematic and taxonomic influence of of the Freshwater Mussels Central study of Frierson. This is understandable, as he New York which Ortmann's complemented was a compiler, not a specialist in the north. This work was followed work to the Unionaeea, and Frierson's work was the the Freshwater Mussels Nova Scotia by of latest synthesis, and should have been the by Athearn and Clarke (1962). last word. This brief review of the litera- primary In 1942, 1949, and 1964, Modell, in ture on the Unionaeea of the Apalachieolan Das natiirliche System der Najaden, pro- and Atlantic itself Slope regions explains posed that the relationships within the the reason for carrying the present study Unionaeea can be found in the beak to the Potomac River. To have carried it sculpture. He proposed forty-three new further would have been redundant, in names for family groupings. He expected the work of Ort- view of the quality of his system to be accepted on the basis of a mann, Clarke and Berg, and Athearn and few diagrammatic sketches of beak sculp- Clarke. ture, and faith in a single character, which I have not found constant. Regarding the Classification of the Unionaeea North American Unionaeea he says: The classification based on the anatomy "... after Simpson's divisions were not of the genera of Unionaeea proposed by sufficient for these purposes [i.e. questions nomenclature]. I had to go along Ortmann (1911, 1912a) is used here. Since concerning completely with the uniform system of Frier- Ortmann's time a few necessary nomen- son in order to give a better picture. With but no clatural changes have been made, this however I also had to use Frierson's 1 one has substantially modified his concepts concepts of Rafinesque's names completely." of the evolution of the North American Of course, as mentioned before, the genera. His objection is to Frierson's systematics. In 1927 Frierson, an ardent naturalist nomenclatural eccentricities, in the sense and an oldfashioned typologist, published implied by Modell, are not of concern here. A Classified and Annotated Check List of

stated 1 the North American Naiades. He Translation by Stansbery and Soehngen, 1964, his method and philosophy: Sterkiana, no. 14, p. 18. 6 296 Bulletin Museum of Comparative Zoology, Vol. 140, No.

of these In short, it is not implied that Ortmanns shall located most figured speci- it in the United States National Mu- anatomical system is perfect, only that mens and has not been improved on. Haas (1969a), seum. These have been rechecked, not use their localities restricted when in SuperfarrriUa Unionacea, does possible. but a mentioned Modell's system of classification for the With few exceptions, all localities were and are higher categories. His species concepts of type relocated, the North American Unionacea are those of often rendered more specific from data a few additions. In on labels and references to Frierson ( 1927 ) , with original by standard modern or Superfamily Unionacea Haas (1969b) only atlases, county maps, United States 1: 250,000 discusses the genera. Geological Survey maps. These additional data are placed in SYSTEMATIC SECTION square brackets. Unless specifically mentioned to the con- The following abbreviations have been trary, all extant types have been examined used in the text and on the plate captions. and usually photographed. Almost none Sciences of ANSP—Academy of Natural of Say's primary types have survived, and Philadelphia, Pennsylvania many of Conrad's are missing, whereas MCZ—Museum of Comparative Zoology, most of Lea's have been located. Cambridge, Massachusetts Only pertinent references are included. MZUM—Museum of Zoology, University Simpson (1914) is referred to only when of Michigan, Ann Arbor, Michigan our species concepts are similar or when USNM—United States National Mu- the differences are easily reconcilable. The Clench and seum, Washington, D. C. papers of Ortmann (1919), Turner (1956), Clarke and Berg (1959). ease of full Synonymy: For reference, all re- and Atheam and Clarke ( 1962 ) are citations are included for each taxon. Else- ferred to when relevant. where in the text, references are abbrevi- Descriptions: The measurements arc ated and require the use of the bibliog- only intended to convey the general size raphy. of specimens from a given station, or to Isaac Lea often rushed brief Latin de- illustrate sexual differences when they are scriptions of his new species into print, a relevant. practice common in his time. These were Anatomy and Breeding Season: These subsequently followed by adequate de- entries are included when such information scriptions and figures, which were then is available. For widely distributed species, reprinted as "Observations on the Genus it should be borne in mind that the dates Unto." Only page references to this work of the season are not necessarily are included here since the plates and breeding applicable to their entire range. figures were never renumbered, but always : Given when known: kept the same numbers as in the preceed- Remarks: Statements made in this sec- ing reference. Lea generally gave several tion, applicable to each species over localities where each of his species had though its entire are designed to been found and did not select types, range, primarily aid in differentiating one species from an- though he always figured a single speci- other within the and At- men, for which he gave measurements. In Apalachicolan lantic lieu of the use of the word type,' under Ar- Slope regions. The are intended to be ticle 73 (b) Int. Code Zool. Nomen. (1964), Range: ranges all-inclusive for each this is an "equivalent expression" and these species. the area specimens are regarded as holotypes. Dur- Records: The records, limited to ing the early part of the century, W. B. Mar- of the study, are based mostly on speci- Atlantic Slope Unionidae • Johnson 297

mens in the major collections mentioned used with specimens primarily from this area. The in Clarke and L959: be used above under Abbreviations. All of these key Berg ( 13) may for the identification of die northeastern have been examined, with a lew species specimens of Unionacea. The key is artificial and is based similar records noted exceptions. Sometimes on shell characters only. Some species of Unioni- arc found in several museums. If so, those dae are quite variable and some specimens may in the Museum of Comparative Zoology not key out correctly. Therefore it is advisable to confirm identifications made with the key by are generally given preference. It is to be reference to the text and figures. assumed that all records are in this 1. Hinge teeth completely absent. 4 museum, unless specifically mentioned to or vestigial hinge teeth the Articulating pres- contrary. ent. 2 A few records from and Virginia Mary- 2. Hinge teeth vestigial. 3 land, of common species, listed by Ort- Articulating pseudocardinal teeth present. 8 mann ( 1919 ) have not been reexamined, 3. Radial wrinkles on the posterior slope, but are accepted with confidence. In each each valve with one thin, depressed tri- of these instances, references are given. angular pseudocardinal tooth; no lateral list but Walter (1956) did not his data, teeth ...Alasmidonta varicosa p. 354 gave numbers on a figure of the Neuse Radial wrinkles absent on the posterior each valve with a River system. Because the first series of slope, slight depression and then a slightly raised thickening just his collection is in the Museum of Com- before the umbos his station numbers parative Zoology, Strophitus undulatus p. 367 follow those of his localities which are 4. Umbos inflated, raised above the hinge line. 5 included here. (See note on p. 323). The Umbos flat, not raised above the hinge records of ubiquitous Elliptio complanata line. 7 have been selected. are more copious They 5. Shell greatly inflated, especially toward is to illustrate the distri- than necessary the posterior ventral margin; Altamaha bution of the species, but they are an River system only „ Anodonta gibbosa p. 359 Shell somewhat but accurate index to the scope of collecting inflated, uniformly so. 6 that has been done in the area covered in 6. Shell uniformly thin, nacre bluish white : this of the portion paper. Anodonto cataracta cataracta p. 356 are In so far as possible, the records Shell prominently thickened along die an- arranged from headwaters to the mouth terior ventral margin below the pallial nacre salmon or colored of the rivers, and from south to north, or line; copper Anodonta implicata p. 360 west to east, as the case may be. Abbeville 7. Dorsal and ventral margins almost parallel; District, South Carolina, which occurs periostracum with broad green rays. Shell many times over in the records, includes very thin and not much inflated 362 the present counties of Abbeville and Anodonta imbecilis p. Greenwood, and most of McCormick. Ventral margin very broadly curved; periostracum with fine green rays. Shell Figures: When available, the holotypes very thin and not much inflated illustrate the various are generally used to Anodonta couperiana p. 365 species. Often more than one illustration 8. Lateral teeth absent or vestigial. 9 lateral teeth 11 is included to show variability. Frequently, Articulating present. 9. Posterior ridge very rounded; shell ellipti- the data on the plate captions is not re- cal or subrhomboid elsewhere. peated _ Alasmidonta nndidata p. 349 Posterior ridge sharp; shell subtriangu- the St. Key to the Unionidae Found Between lar. 10 and the Potomac Marys River, Florida, 10. Umbos anterior to the center, incurved; Maryland. River, posterior slope acute Alasmidonta 351 The following key includes all of the species of triangulata p. Unioniclae that are found between the St. Marys Umbos centrally located, acutely incurved; at almost 90° angle with and the Potomac rivers, and it is designed to be posterior slope 298 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

posterior ridge; Altamaha River system descending posterior margin, rendering only Alasmidonta areola p. 352 the posterior end rather wedge shaped. 22 11. Lateral teeth double in the right valve, 21. Shell rather uniformly trapezoidal, disk single in the left one. Shell quite small flattened; periostracum not usually shiny, Alasmidonta heterodon p. 347 often rayed, yellowish green to black Lateral teeth single in the right valve, Elliptio complanata p. 314 double in the left one. 12 Shell subrhomboidal, often somewhat 12. Pseudoeardinal teeth chunky and serrated, pointed, very variable as to shape and the more prominent one in the right valve degree of inflation; periostracum usually not parallel to the hinge line. 13 subshiny to shiny, often rayed, yellowish Pseudoeardinal teeth long and lamellate, to brownish . ...Elliptio icterina p. 325 with a tendency of the larger one in the 22. Valves with greatest degree of inflation right valve to be parallel to the hinge at midpoint of posterior ridge; shell often line. 29 rayed when young, becoming blackish 13. Shell less than, or about, twice as long as widi age. Periostracum smooth on the high. 14 disk, rough toward the margins; Altamaha Shell distinctly more than twice as long River system only Elliptio hopctonensis p. 324 as high. 37 Valves rather uniformily inflated; shell not 14. Posterior ridge with faint ridges crossing rayed, occasionally greenish when young, lines of growth. 15 becoming blackish with age. Periostracum Posterior ridge without faint ridges cross- generally distinctly satiny ing line of growth. 20 Uniomerus tetralasmus p. 339 15. Shell subrhomboidal, subtriangular or 23. Shell small, not exceeding 65 mm in quadrate, often solid, sometimes quite length, seldom rayed; periostracum yel- ponderous. 16 lowish, greenish, or brownish. Sexual Shell subelliptical or elongated rhomboid, dimorphism generally present. 24 generally quite thin. 19 Shell large, generally rayed, periostracum 16. Posterior ridge of variable acuteness. 17 greenish yellow. Sexual dimorphism pres- Posterior ridge sharp; shell rather thin, ent. 28 widi green rays; Altamaha River system 24. Shell without spines on disk. 25

only . ..Elliptio dariensis p. 310 Shell with spines on disk. Outline sub- 17. Shell ponderous, not rayed, or with thin quadrate or subrhomboid; periostracum dark green rays when young, generally yellowish or brownish. Sexual differences blackish; St. Marys River system only not well marked Pleurobema collina p. 300 Elliptio c. crassidens p. 305 25. Shell with a constriction of the ventral Shell not ponderous, with green rays of margin before a double, but faint, pos- varying width, yellowish or chestnut terior ridge; periostracum smooth. 26 .. p. 308 Shell with a regularly rounded ventral 18. Posterior ridge sharp near the umbos, be- margin before a sharp double posterior coming faint and biangulate toward the ridge; periostracum with numerous heavy posterior margin. Shell compressed, brown- ridges. Outline of female long obovate; ish. Young specimens occasionally with outline of male elliptical faint green rays Elliptio fratema p. 312 Carunculina pulla p. 370 Posterior ridge consistently acute. Shell 26. Outline subquadrate or rhomboid; perios- subinflated, brownish, often with narrow tracum often clothlike, blackish brown,

green rays over entire surface - occasionally with rays. Sexual difference ..Elliptio ivaccammuensis p. 313 not well marked ...Pleurobema masoni p. 301 19. Shell ponderous, inflated; posterior ridge Outline of female short obovate; of male double and of variable acuteness, brown- subelliptical. Periostracum yellowish green, ish, rhomboid to obovate; Satilla River widi fine green rays over the entire sur-

system only . Elliptio c. downiei p. 307 face — . —Vittosa constricta p. 378 Shell not ponderous, disks flattened; pos- 27. Outline a long ellipse; greenish rays over terior ridge double, but uniformly entire surface, periostracum satiny and

rounded. 20 clothlike - Villosa villosa p. 372 20. Dorsal margin long, forming an acute Shell not as above and much larger. 28 angle with the obliquely descending pos- 28. Posterior ridge well developed, especially terior margin. 21 toward the umbos. Shell subovate with Dorsal margin very long, forming an ex- narrow or wide rays generally distributed tremely acute angle with the obliquely over a rather shiny yellowish background. Atlantic Si oil Unionidae • John-son 299

Sexual differences well marked in the ventral sulcus. Dorsal and ventral margins shell - Lampsilis ovata p. 386 roughly parallel. 39 Posterior ridge low and rounded. Shell 38. Shell somewhat over twice as long as suhelliptical, with wide rays generally high, suhelliptical, posterior end pro- distributed over a dull yellowish green duced; seldom rayed Elliptio arctata p. 331 background. Sexual differences not al- Shell over three times as long as high, ways well marked in the shell trapezoidal, with the highest point at the Lampsilis radiata radiata p. 390 anterior end tapering to a long point; 29. Left valve without a rudimentary inter- rays visible when young. Altamaha River dental projection. 30 only Elliptio shepardiana p. 338 Left valve with a rudimentary projection, 39. Nacre silvery white; posterior end dis- which fits into an interdental groove in tinctly lance-head shaped; rays always the right valve; shell generally small, 45 present. Sexual dimorphism present. Fe- to 65 mm in length, though giant speci- males with a distinct postbasal swelling. mens occur; subrhomboid, often with a Ligumia nasuta p. 380 low posterior wing when young . Nacre coppery or purple; dorsal and vent- ...Lasmigona suhoiridis p. 343 ral margin almost parallel, often rayed, over twice as as 30. Shell without spines on surface. 31 somewhat long high laneeolata 333 Shell with hollow spines in front of a Elliptio p. sharp posterior ridge. Outline subtri- nacre Altamaha River 1 angular; purple. Superfamily Unionacea Thiele 1935 system only p. 303 31. Posterior ridge present and well defined, Family Unionidae (Fleming 1828) Ortmann at least toward the umbos. Shell rather 1911 large and solid, yellowish, with varying amounts of the surface covered with Subfamily Unioninae (Swainson 1840) green rays. Sexual dimorphism present. 32 Ortmann 1910 Posterior ridge very broadly rounded. Genus Pleurobema Shell small and thin, with numerous green Rafinesque rays over the entire surface, more greenish Subgenus Pleurobema s.s. Rafinesque than yellow. 35 32. Surface of the shell either without rays Pleurobema Rafinesque 1819, Jour. Phys. Chim. or with some on posterior slope only. Hist. Nat. (Paris), 88: 427. Species listed: P. Periostracum shiny. 33 mytiloides Rafinesque, P. conica ? Rafinesque Surface of the shell with rays over the [nomina nuda]. Rafinesque, 1820, Ann. Gen. entire surface. Periostracum more often des Sci. Phys. (Bruxelles), 5: 313. Species subshiny than shiny. 34 listed: P. mytiloides Rafinesque, P. cuneata 33. Posterior ridge low and rounded except Rafinesque. interdentum toward the umbos, present. Type species, Unio mytiloides Rafinesque. Sub- 382 Nacre bluish white ..Lampsilis cariosa p. sequent designation, Herrmannsen, 1847, Indi- Posterior ridge very sharp, interdentum cis Generum Malacozoorum, 1: 292. present; Altamaha River system only . Ortmann, 1912, Ann. Carnegie Mus., 8: 261. Lampsilis dolabraeformis p. 384 Ortmann and Walker (1922, Occ. Pap. Mus. 34. Nacre bluish; rays narrow. Posterior ridge Zool., Univ. Mich. no. 112, p. 20) state that rounded except toward the umbos. Shell the description of P. mytiloides in 1820 ren- thin, no interdentum dered the genus monotypic. This is not so, as Lampsilis ochracea p. 388 the first use of P. mytiloides has no bearing on 35. Periostracum shiny. 36 the subsequent recognition of the genus. Periostracum distinctly satiny. s. Frierson Villosa villosa p. 372 Under Pleurobema s., (1927: and 36. Narrow green rays, interrupted by growth 40_44) includes thirty-seven species rests Villosa delumhis 375 of p. subspecies. It is probable that some Wide not by green rays, interrupted these taxa will be subsequently reduced to growth rests Villosa vibcx p. 373 but it is clear that this genus 37. Shell with ventral margin incurved, often synonymy, sufficiently as to render entire shell ar- 1 is the standard cuate. 38 Or, Unionoidea ( oidea ending acea has been Shell with ventral margin straight or for superfamily names, though gen- used in molluscan taxonomy). curved, sometimes with a slight dorsal- erally 300 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

has achieved its greatest diversity in the larly rounded; posterior end angular and Interior Basin. slightly cut away below. Ventral margin straight or slightly arcuate centrally. Dor- Subgenus Lexingtonia Ortmann sal margin almost straight, terminating in a Lexingtonia Ortmann 1914, Nautilus, 28: 28. sharp angle with the obliquely truncated Type species, Unio subplanus Conrad. Original posterior margin. Hinge ligament short but designation. prominent. Posterior ridge full, rounded, toward the base of the Ortmann instituted Lexingtonia as a weakly biangulate shell. Umbos elevated and located genus because he believed that the six to slightly somewhat anterior to the middle of the eight subconcentric ridges of the beak shell, their of three to sculpture and the red placentae differenti- sculpture consisting four subconcentric that form ated it from Elliptic- and Pleurobema. He strong ridges mentions that the shell resembles Fusconaia an indistinct rounded angle on the pos- terior Surface of the disk in which all four gills serve as marsupia, ridge. smooth, with undifferentiated concentric but since he also says that only the outer growth = rings, surface with short differ- gills are so used by U. sul)planus Conrad sculptured masoni Conrad, as in Pleurobema, the entiated bilateral spines especially in the of the dorsal and present author is placing Lexingtonia under region posterior slope of the disk. Periostracum Pleurobema as a subgenus, as did Frierson posterior quarter or in (1927: 44). bright yellow greenish yellow young brownish in Under Lexingtonia, Frierson (1927: 44, shells, becoming adults; rarely with brownish 45) includes three species. Two of these rayed fine, narrow, broken, are included here under the synonymy of lines. Pleurobema (Lexingtonia) masoni (Con- Left valve with two ragged, triangular rad). The third, Pleurobema utriculum pseudocardinal teeth and two short, (Lea) belongs to the Tennessee River obliquely descending parallel lateral teeth. drainage. Its synonymy has not been re- Right valve with one ragged, triangular investigated, but it probably does not be- pseudocardinal with a vestigial tooth above long in this subgenus, which appears to it; one lateral tooth. Beak cavities com- be restricted to the Southern Atlantic pressed; anterior adductor muscle scars Slope. deep, posterior ones very faint. Pallial line distinct, especially anteriorly. Nacre white, Pleurobema (Lexingtonia) collina (Conrad) occasionally slightly pinkish, often bluish Plate 2: 1-2 posteriorly. Unio collinus Conrad 1836, Monography Unioni- Length Height Width dae, no. 7, pi. 36, fig. 2 [plate caption only]. mm mm mm 1837, op. cit., no. 8, p. 65 (North [= Maury] 55.0 34.7 20.3 Rivanna River, 2 mi. W River, a branch of the River [Rockbridge James Columbia, Fluvanna Co., Co.], Virginia; type lot ANSP 20408 [lost]). Virginia. 1840, op. cit., no. 12, p. 109, pi. 60, fig. 3; 46.0 30.0 19.0 North River [Rockbridge this subsequently figured metatype, here se- Co.], Virginia. Lecto- lected, lectotype ANSP 41007. type. Alasmidonta collina (Conrad). Simpson, 1914, Cat. Naiades, 1: 501. Anatomtj. Discussed by Boss and Clench Pleurobema collina (Conrad). Boss and Clench, (1967: 47). 1967, Occ. Papers on Moll., 3: 45-51. Breeding season. Probably tachytictic, Description. Shell small, seldom reach- releasing the glochidia during the sum- ing more than 55 mm in length. Outline mer. (Boss and Clench, 1967: 48.) subquadrate or subrhomboid. Valves sub- Habitat. Lives on sandy bottoms, in compressed, subsolid. Anterior end regu- rather swift water. • Atlantic Sloi i Unionidae Johnson 30]

Remarks. Pleurobema collina (Conrad) Pleurobema (Lexingtonia) masoni (Conrad) is a distinct Southern Atlantic Slope Plate 2: 3-10 species. It is remarkable for the spines on Unio inaso)ii Conrad 1834, New Fresh Water the posterior slope of the valves. Usually Shells United States, p. 34, pi. 5, fig. 2 the maximum number of pairs is three, (Savannah River, Augusta [Richmond Co.], in are and, general, they nearly bilaterally Georgia; figured holotype ANSP 41333). Con- symmetrical. The first set of spines is about rad, 1836, Monography Unionidae, no. 3, p. 3-4 mm from the umbo, the second 6-8 28, pi. 12, fig. 2; figured paratype ANSP 41332. Unio subplanus Conrad I Unioni- and the third from 10-17 mm. The

seum ) . Description. Shell small, seldom reach- River System James ing more than 50 mm in length. Outline Valves sub- James River Drainage. Virginia: Calf- subquadrate or rhomboid. pasture River (Conrad, 1846: 407); North compressed, subsolid. Anterior end regu- [= Maury] River, Lexington; James River, larly rounded; posterior end angular and near Natural Bridge (USNM), all Rock- slightly cut away below. Ventral margin bridge Co. James River, Buchanan, Bote- straight save for a slight undulation pos- tourt Co. Rivanna River, 2 mi. W teriorly. Dorsal margin straight, terminat- Columbia, Fluvanna Co. James River, op- ing in a sharp angle with the obliquely posite Maidens, Goochland Co. truncated posterior margin. Hinge ligament 302 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

short but prominent. Posterior ridge full, Clench and Turner, 1956: 162, pi. 8, fig. rounded, often with a second or tertiary 6) which ranges from the Apalachicola to ridge ending in slight biangulations toward the Suwannee River system. The latter the base of the shell. Umbos slightly ele- species is decidedly more subovate, with vated, located somewhat anterior to the the posterior end tending to be broadly middle of the shell, their sculpture con- pointed. The ventral margin is rarely sisting of six to eight rather crowded sub- arcuate posteriorly; the disk is less com- concentric ridges that form an indistinct pressed. The periostracum is often blackish rounded angle on the posterior ridge, in or yellowish brown, but seldom rayed. front of which they are wavy and corru- Except for examples of P. masoni from gated. Disk often with irregular incre- the Nottoway and James rivers, near the mental striae, which sometimes render the northern limit of its range, where the shells periostracum rather clothlike; greenish, reach their maximum size and are found in yellowish, or yellowish brown, sometimes relative abundance, this is a very rare with dark, dull greenish rays. species. In most drainage systems it is Left valve with two ragged, triangular known from but a few examples. In 1961, pseudocardial teeth and two short, oblique Clench, Boss, and Fuller collected several descending, parallel lateral teeth. Right thousand LTnionidae in Georgia, but found valve with one ragged, triangular pseudo- only seven specimens of this species from cardial with a vestigial tooth above it; one two localities. lateral tooth, which is sometimes double. In the James River, Virginia, P. masoni Beak cavities compressed; anterior adduc- is found closely associated with P. eollina, tor muscle scars deep, posterior ones very under which see: Remarks on p. 301. faint. Pallial line distinct anteriorly only. Range. Southern Atlantic Slope: Nacre white or bluish white, thinner and Ogeechee River system, Georgia, north to somewhat irridescent posteriorly. the James River system, Virginia.

Width Lengthmm Heightmm mm Specimens Examined 62 41 22 3 mi. E Nottoway River, Ogeechee River System Rawlings, Brunswick Co., Virginia. Ogeechee River Drainage. Georgia: 47 28 19 Mill Race, 2 mi. N Sardis, Ogeechee River, 12 mi. SW Warrington, Bnrke Co., Georgia. Warren Co. Buckhead Creek, 10 mi. SW 31 21 14 12 Ogeechee River, Shoals, Sardis, Burke Co. mi. S Warrington, Warren Co., Georgia. Savannah River System Savannah River South Anatomy. Discussed by Ortmann, 1914, Drainage. Caro- lina: Nautilus, 28: 29-30. "Glochidia semi- Abbeville District (USNM). Georgia: Broad River or Elbert Co.] elliptical, without hooks. Length and [Madison Savannah Rich- height the same, 0.18 mm." (MZUM). River, Augusta, mond Co. Mill Race, 2 mi. N Breeding season. Ortmann found a (ANSP). Sardis, Burke Co. gravid female in the North River, Lexing- ton, Rockbridge Co., Virginia, on June 7, Cooper-Santee River System 1912. Catawba River Drainage. North Caro- Habitat. Lives on bottoms, in sandy lina: Long Creek, Gaston Co. rather swift water. Remarks-. Pleurobema masoni (Conrad) Pedee River System is a distinct Atlantic Slope species. It is Yadkin River Drainage. North Caro- closest to P. pyriforme (Lea) (figured by lina: Salem, Forsyth Co. (MZUM). Atlantic Slope Unionidae • Johnson 303

Cape Fear River System erection of a separate genus: Canthyria" for this 2: Cape Fear River (MZUM). species. Simpson (1914, 704) states that the anatomy of spinosa is typical Neuse River System of Unio [=Elliptio]. Frierson (1927: 50) used this name as a to include both Neuse River Drainage. North Carolina: genus and Pleurobema collina Hillsboro, Orange Co. Eno River, 6 mi. spinosa (Conrad) on the basis of the Ross and Clench NNW Durham Center, Durham Co. (112). spines. have shown that these Walnut Creek, Raleigh (USNM); Neuse (1967: 50) clearly two are of different and River, Raleigh; Little River, 2 mi. WSW species lineages, that the are a character. Zebulon (6); all Wake Co. Rlack Creek spines convergent In of the anatomical to [about 10 mi. W Renson, Johnston Co.] spite similarity the shell is not close (USNM). Elliptio, morphology to that of any other member of the genus. Pamlico River System This appears to be a very old species, which is restricted to the Altamaha River Tar River Drainage. North Carolina: Tar River, Providence, Granville Co. Tar system, Georgia. This is River, Rruce, 9 mi. NW Greenville; Chicod subgenus monotypic. Creek, 8 mi. W Greenville; both Pitt Co. Elliptio {Canthyria) spinosa (Lea) Roanoke River System Plate 2: 11 River North Caro- Roanoke Drainage. Unio spinosus Lea 1836, Description of a new lina: Roanoke River, Weldon, Halifax Co. Unio, pp. 1-4 (col. figs.) [not seen]. Lea, 1838, Trans. Amer. Philos. Soc, 6: 57, pi. 16, fig. Chowan River System 50 (Altamaha [River], Hopeton, near Darien [Mcintosh Co.], Georgia; figured type in Nottoway River Drainage. Virginia: collection of Prof. Shepard [presumed lost]; 3 mi. E Rruns- Nottoway River, Rawlings, Altamaha [River], Liherty [now Long] Co., wick Co. Georgia). Lea, 1838, Obs. Unio, 2: 57. Simp- son, 1914, Cat. Naiades, 2: 702. James River System Description. Shell medium to large, James River Drainage. Virginia: Calf- reaching 110 mm in length. Outline pasture River (Conrad, 1846: 407); North subrhomboidal or subtriangular. Valves [=Maury] River, Lexington (ANSP); North subinflated to inflated, inequilateral, or River, above Ruena Vista (Carnegie Mus.); almost equilateral; shell solid. Anterior end [North River] Natural Rridge; all Rock- regularly rounded; posterior end slightly bridge Co. Rivanna River, 2 mi. W Co- broader, ending in a point just below the lumbia, Fluvanna Co. James River, oppo- medial line. Ventral margin slightly curved. site Maidens, Goochland Co. James River, Dorsal margin obliquely descending from Richmond, Henrico Co. (MZUM). the umbos, imperceptibly joining the de- scending posterior margin. Hinge ligament Genus Elliptio Rafinesque prominent, but short. Posterior ridge Subgenus Canthyria Swainson sharply angular, often with a faint second- which renders the Canthyria Swainson 1840, Treatise on Malacology, ary ridge above it, Umbos pp. 276, 378. posterior slope slightly rugose. Unto Lea. Original desig- the lo- Type species, spinosus slightly elevated above hinge line, nation. cated in the anterior third of the shell, their of several Ortmann (1912, Ann. Carnegie Mus., 8: sculpture consisting very tiny of the shell smooth 269) suggests that "the spines [of the type] bars. Surface generally with fine concentric are unique, and would possibly justify the and shiny sculpture. 304 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Each valve ornamented with from one to a height of about 30 mm. Presumably it five spines of various length (varying from is while the is young that the spines 1-2.5 mm in length) straight or crooked, are most useful as grappling devices for usually arranged in a row, roughly parallel holding the shells in sand. to the posterior ridge. Periostracum green- Pleurobema coUina (Conrad) of the Tar ish yellow or brownish in old shells, usually and James rivers of North Carolina and with faint greenish and yellowish rays. Virginia is also spined. This led Frierson Left valve with two pseudocardinal (1927: 46) to unite the two species under teeth, one in front of the other, the an- the genus Canthyria Swainson 1840, but terior one somewhat triangular, the hinder Boss and Clench (1967: 50) have clearly one vestigial. Hinge line short and narrow, shown that the two species are of different two obliquely descending straight lateral lineages, and that the spines are a con- teeth. Right valve with two roughly vergent character. parallel pseudocardinals, the posterior one The distinct triangular shape; the equally triangular and serrated, the more anterior distinctive greenish yellow, finely rayed, one low and vestigial; one lateral tooth. periostracum; the pink or purplish nacre; Beak cavities moderately deep, with a and the long spines distinguish E. spinosa few dorsal muscle scars. Anterior and from all other Atlantic Slope unionids. posterior adductor muscle scars well im- In his "Travels of William Bartram," the pressed. Pallial line distinct. Nacre pink- well-known botanist mentions seeing this ish or purplish, seldom white. shell in 1773, probably near Frederica The spine is a tubular growth which [Glynn Co., Georgia], but writing his appears to contain a lobe of the mantle journal from memory some thirteen years until it has reached its full height, at which later, he claims to have seen it in the time the lobe fills the hollow spine with Mississippi River, Louisiana (Goodrich, shell as it retreats. 1930: 140). Range. Southern Atlantic re- Length Height Width Slope: mm mm mm stricted to the Altamaha River system, 110 54 48 Oconee River, 2.5 mi. N Georgia. Glenwood, Montgomery Co., Georgia. Specimens Examined 89 54 37 As above. Altamaha River System 69 43 35 Altamaha River, 4 mi. NE Jesup, Wayne Co., Ocmulgee River Drainage. Georgia: Georgia. Ocmulgee River, below Lumber City, Tel- fair Co. (H. D. Athearn). Anatomy. Discussed by Lea (1863: Oconee River Drainage. Georgia: Oco- 413). The anatomy is typical of Undo nee River, 2.5 mi. N Glenwood, Montgom- [=Elli))tio] according to Simpson (1914, ery Co. 2: 704). Altamaha River Drainage. Georgia: Habitat. Lives buried from two to four Altamaha River, 10 mi. ENE Surrency, inches deep in sand bars, in swift water. Appling Co. Altamaha River, 11 mi. N Remarks. EUiptio spinosa (Lea) is Odum; Altamaha River, 4 mi. NE Jesup; known only from the Altamaha River both Wayne Co. Altamaha River, Fort system of Georgia. It is remarkable for the Barrington; Altamaha River, Hopeton, near long spines that run to the parallel pos- Darien; both Mcintosh Co. terior ridge that are erected near growth rests the first few None of during years. Subgenus EUiptio s.s. Rafinesque the some sixty specimens seen have spines EUiptio Rafinesqne 1819, Jour. Phys. Chim. Hist. that are started after the shells had reached Nat. (Paris), 88: 426 [nomen nudum]. Rafi- Atlantic Slope Unionidae • Johnson 305

nesque, 1820, Ann. Gen. Sci. Phys. (Bruxelles), the species of EUiptio s. s. are the most 5: 291. listed: E. E. Species nigra Rafinesque, abundant Unionidae. EUiptios are often crassa (Say), E. oiridis Rafinesque, E. fasciata found in environments in which no other Rafinesque. genera of Unionidae live, since some of them Type species, Unto nigra Rafinesque. Subsequent designation, Ortmann, 1912, Ann. Carnegie have an unusually wide environmental Mas., 8: 266. The previous use of Unio crassi- tolerance, even to silting and pollution. dens Lamarck as tvpe species by Simpson As a consequence, some of the species have Proc. U. S. 'Natl. 22: 700) is (1900, Mus., and at a invalid since Lamarck's name was not included developed many ecophenotypes, station there often to be less by Rafinesque in his list of species. given appears variation than Eurynia Rafinesque 1819, Jour. Phys. Chim. Hist. interspecific infraspecific Nat. (Paris), 88: 426 [nomen nudum]. Rafi- variation between localities. This variation Ann. Gen. Sci. nesque, 1820, Phys. (Bruxelles), has led to a plethora of names being 5: 297. Species listed: Unio dilatata. Unio applied to the several species. latissima, Unio solenoides, all Rafinesque. Type species, Unio dilatatus [sic] Rafinesque. crassidens crassidens Subsequent designation, Herrmannsen, 1847, EUiptio (EUiptio) Indicis Generum Malacozoorum, 1: 436. (Lamarck) Cunicula Swainson 1840, Treatise on Malacology, Plate 3: 1-3 pp. 268, 378. Species listed: C. planulata Lea, Unio crassidens var. b Lamarck 1819, Hist. Nat. C. patula Lea, C. rubiginosa Lea, C. secura des Animaux sans Vertebres, 6: 71 (lac Erie Lea, C. purpurascens Lamarck. [erroneous]; lectotype, Paris Museum, selected Type species, Unio purpurascens Lamarck. Sub- by Johnson, 1969, Nautilus, 83: 53, fig. 2, and sequent designation, Herrmannsen, 1847, In- the type locality restricted to the Ohio River, dicis Generum Malacozoorum, 1: 335. Cincinnati, Ohio). A composite species, re- Ortmann and Walker (1922, Occ. Pap. Mus. Philos. stricted by Lea ( 1834, Proc. Amer. Soc, Zool., Univ. Mich. no. 112, p. 28) selected 5: 87). as first revisor, on the basis of his EUiptio over Eurynia on the basis of page prece- identification of the = Unio cuneatus dence. type Barnes. See: Ortmann and Walker (1922: 28). Unio Ann. Gen. Sci. Although EUiptio has been usually nigra Rafinesque 1920, Phys. 5: 1-4 (Ohio treated as a masculine noun, H. B. Baker (Bruxelles), 291, pi. 80, figs. River; type ANSP 20243, teste, Vannetta, 1915, 78: has out (1964, Nautilus, 33) pointed Proc. Acad. Nat. Sci. Phila., 66: 555). that Bafinesque consistently used EUiptio Unio cuneatus Barnes 1823, Amer. Jour. Sci., 6: as feminine (e.g. E. nigra) and therefore 263 (Ohio River; type [lost]). the name should be thus treated. Unio incrassatus Lea 1840, Proc. Amer. Philos. Soc, 1: 268 (Chattahoochee River, near Co- EUiptio s. s. is represented by EUiptio lumbus [Muscogee Co.], Georgia). Lea, 1840, crassidens (Lamarck) and E. dilatata Trans. Amer. Philos. Soc, 8: 217, pi. 16, fig. 34; 1919: (Bafinesque), (figured by: Ortmann, figured holotype USNM 84537). Lea, 1842, 95, pi. 8, fig. 2) in the Interior Basin, Obs. Unio, 3: 55. Conch. where they are widespread and locally Unio discus Sowerby 1868, in Reeve, Iconica 16, Unio, pi. 62, fig. 310 (India) non abundant. At present, it is a moot question Lea 1838. whether the several subgenera from Central Unio lehmanii S. H. Wright 1897, Nautilus, 10: included Frierson 33- America, by (1927: 138 (St. Marys River, [Nassau Co.], Florida; 39) under EUiptio, belong there, but if lectotype USNM 149650 selected by Johnson, Occ. 3: 3). they do, the evidence is that the area com- 1967, Pap. Moll., 7, pi. 6, fig. Unio daniehii B. H. Wright 1899, Nautilus, 13: prised of Mexico and Central America is 31 (Spring Creek [a branch of the Flint River], one where considerable has oc- speciation Decatur Co., Georgia; measured holotype curred in this genus. USNM 168967, figured by Johnson, 1967, Occ. 3: Speciation within EUiptio s. s. has oc- Pap. Moll., 5, pi. 6, fig. 1). Unio polymorphic B. H. Wright 1899, Nautilus, curred primarily in the Apalachicolan 13: 42 (Spanish Creek [a tributary of the St. Peninsular and the South- region, Florida, Marys River, W of Folkston], Charleton Co., ern Atlantic Slope region. In these regions Georgia; lectotype USNM 152060 selected by 306 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Johnson, 1967, Occ. Pap. Moll., 3: 8, pi. 6, fig. generally uniformly black. Left valve with 2). two heavy, rough pseudocardinal teeth, the Elliptic pachyodon Pilsbry 1953, Pliocene Moll. more anterior one smaller and lower. Southern Florida, Acad. Nat. Sci. Phila., Monog. Hinge line rather short and 8, p. 447, pi. 65, fig. 8 (St. Petersburg [Hills- broad, with two short, boro Co.], Florida; type ANSP 18586; Plio- slightly curved lateral teeth. Right valve cene). with one chunky serrated pseudocardinal; Elliptio niger Rafinesque. Ortmann 1919, Mem. one lateral tooth. Beak cavities very shal- Carnegie Mus., 8: 91, pi. 8, fig. 1. low, with a few dorsal muscle scars under incrassatus (Lea). Clench and Turner, 1956, Bull. Florida State Mus., 1: 171, the hinge plate. Anterior adductor muscle pi. 8, fig. 1. scars very deep, posterior ones and pallial line distinct. Nacre usually purplish, Description. Shell often large, exceed- salmon, sometimes yellowish, posteriorly ing 130 mm in length, though usually not iridescent. exceeding 100 mm in the Atlantic drain- age. Outline subrhomboid, or quadrate, Lengthmm somewhat produced posteriorly in older specimens. Valves somewhat inflated, rather solid to ponderous, inequilateral. Anterior end regularly rounded; posterior end more broadly rounded and either somewhat pointed or biangulate. Ventral margin straight or slightly curved, some- times arcuate in matures. Dorsal margin broadly rounded, usually indistinctly join- ing the obliquely descending posterior margin. Hinge ligament prominent, lo- cated posteriorly. Posterior ridge generally sharp and angular, especially when young, often becoming more broadly rounded with age. Posterior ridge generally double. Secondary ridge varying greatly as to prominence and distance from the primary ridge. This variation causes a considerable range as to degree of terminal biangulation, and further causes the posterior slope to vary from subtruncated to broad. Posterior slope generally with a few wrinkles radiat- ing from posterior ridge to dorsal margin. Umbos rather full, but rather low, located in anterior third of shell, their sculpture consisting of a few coarse ridges running parallel to growth lines. Young shells may be rather smooth on the disk, developing concentric striae with age; posterior slope usually becoming more wrinkled. Perios- tracum fine when young, reddish brown or chestnut, sometimes yellowish with greenish rays; posterior slope generally darker. In older specimens, periostracum Atlantic Slope Unionidae • Johnson 307

concomitant posterior biangulation, and the Description. Shell large, reaching up to breadth or acuteness of the posterior slope. 100 mm in length. Outline obovate, sub- There is a general tendency for the poste- obovate, or rhomboid. Valves rather in- rior ridge to be rounded and the posterior flated, solid, inequilateral. Anterior end slope to be broad in specimens from the regularly rounded; posterior end broadly Apalachicolan region. rounded, slightly biangulate, or truncate. E. c. crassidens can be confused with Ventral margin straight or curved, some- some allopatric species in the Southern times rendered arcuate by slight postbasal Atlantic Slope region, but in general, the swelling. Dorsal margin broadly rounded, thickness of the shell, the large, rough, forming a slight angle, occasionally a very pseudocardinals, and short hinge line, dis- sharp one, with the obliquely descending tinguish it from other species of Elliptic posterior margin. Posterior ridge varies Range. Interior Basin: Mississippi drain- from being rather sharp to almost imper- age generally. West Gulf Coastal region, ceptible. Secondary ridge usually present. Alabama-Coosa River system, and Apalach- Posterior slope subtruncated to broad. icolan region: Amite River, Louisiana, Umbos rather full, but low, located in an- east to the St. Marys River system, Florida. terior fifth of shell, their sculpture con- Lea in the of a few coarse Replaced by Elliptio fraterna ( ) sisting ridges running Choctawhatchee River system and missing parallel to growth lines. Periostracum fine from the Ochlockonee and Suwannee river when young, yellowish or brownish with or chest- systems. Peninsular Florida: St. Peters- green rays, becoming dull brown

Pliocene . nut with varies from clothlike burg, Hillsborough Co. ( ) age, being to smooth and shiny. Specimens Examined Left valve with two heavy, rough pseudocardinal teeth, the more anterior St. Marys River System one somewhat triangular. Hinge line short St. River Marys Drainage. Georgia: and broad, with two short, slightly curved 2 mi. E St. St. Marys River, George; lateral teeth. Right valve with one chunky, St. Spanish Creek, W Folkston; Marys serrated pseudocardinal; one lateral tooth. 4 mi. SSE all Charleton River, Folkston; Beak cavities very shallow, with a few Co. dorsal muscle scars under the hinge plate. Anterior adductor muscle scars very deep, crassidens downiei (Lea) Elliptio {Elliptio) posterior ones and pallial line distinct. Plate 4: 1-4 Nacre purplish or flesh-colored, rather dull. Unto downiei Lea 1858, Proc. Acad. Nat. Sci. Length Height Width Phila., 10: 166 (Buck Lake, a bayou of the mm mm mm Satilla River, Wayne [Brantly] Co., Georgia). 95 57 42 Satilla River, 3 mi. S Hor- Lea, 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: Geor- USNM tense, Brantly Co., 210, pi. 25, fig. 91; figured holotype gia. 84854. Obs. Unio, 7: 28. Lea, 1859, 84 49 36 Buck Lake, a bayou of the Unio satillaensis Lea Proc. Acad. Nat. Sci. 1858, Satilla River, Brantly Co., 10: 166 Camden Co., Phila., (Satilla River, Georgia. Holotype of U. Georgia). Lea, 1859, Jour. Acad. Nat. Sci. downiei Lea. Phila., ser. 2, 4: 216, pi. 27, fig. 96; figured 76 57 40 Satilla River, Wayne Co., holotvpe USNM 84855. Lea, 1859, Obs. Unio, Georgia. Holotype of U. 7: 34. spissus Lea. Unio spissus Lea 1859, Proc. Acad. Nat. Sci. Phila., 11: 112 (Satilla River, Wayne Co., Habitat. Lives in sand in swift water. Georgia). Lea, 1859, Jour. Acad. Nat. Sci. Remarks. Elliptio crassidens downiei Phila., ser. 2, 4: 208, pi. 25, fig. 89; figured holo- is quite distinct from the highly type USNM 84853. Lea, 1859, Obs. Unio, 7: (Lea) a of 26. variable typical form, and has pattern 308 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

variability of its own, even though oc- Unionidae, no. 11 [back cover] (Black Water Neuse Yadkin casional individuals are found that are very River, Virginia; River; River; both North Carolina). 1840, op. cit., no. 12, p. close to some specimens from the nearby 103, pi. 57, fig. 1; type from Yadkin River, not St. River in the Marys system. Found only in ANSP [lost]. The specimens from the ad- Satilla River, downiei is usually consider- ditional localities were probably Elliptio com- ably more inflated than the typical form. planata ( Lightfoot ) . Unio Lea Proc. Acad. Nat. Sci. There is a tendency for the shells from pusillus 1840, Phila., 1: 286 (Ogeechee River, Liberty Co., the upper reaches of the river to be Georgia). Lea, 1842, Trans. Amer. Philos. Soc, obovate or with the elongate posterior 8: 220, pi. 18, fig. 36; figured holotype USNM ridge almost imperceptible and the poste- 85241. Lea, 1842, Obs. Unio, 3: 58. rior slope very broad. Some individuals are Unio sordidis Lea 1852, Trans. Amer. Philos. Soc, 10: 254, pi. 12, fig. 1 (Abbeville District also sometimes slightly swollen in the post- [Savannah River drainage], South Carolina; basal In the lower of the river, region. part figured holotype USNM 85688). Lea, 1852, the shells, in general, become more Obs. Unio, 5: 10. rhomboidal, the posterior margin appear- Unio gibbesianus Lea 1852, Trans. Amer. Philos. Soc, 10: 254, pi. 12, fig. 2 District ing truncate with the posterior ridge (Abbeville [Savannah River drainage], South Carolina; and the The angular posterior slope sharp. figured holotype USNM 85685). Lea, 1852, posterior slope of downiei lacks the radial Obs. Unio, 5: 10. wrinkles usually present in E. c. crassidens; Unio rufusculus Lea 1852, Proc. Amer. Philos. 5: 252 District [Savannah the periostracum with its satiny clothlike, Soc, (Abbeville River drainage], South Carolina). Lea, 1852, or even shiny appearance, and yellowish Trans. Amer. Philos. Soc, 10: 258, pi. 14, fig. brown to chestnut color is also distinctive. 7; figured holotype USNM 85683. Lea, 1852, Range. Apalachicolan region: restricted Obs. Unio, 5: 14. to the Satilla River system, Georgia. Unio jorbesianus Lea 1852, Proc. Amer. Philos. Soc, 5: 251 (Savannah River, Georgia). Lea, Specimens Examined 1852, Trans. Amer. Philos. Soc, 10: 264, pi. 16, fig. 17; figured holotype USNM 84542. Satilla River System Lea, 1852, Obs. Unio, 5: 20. Unio buxeus Lea 1852, Trans. Amer. Philos. Soc, Satilla River Satilla Drainage. Georgia: 10: 261, pi. 15, fig. 13 (Abbeville District River, Wayne Co. Buck Lake, a bayou of [Savannah River drainage], South Carolina; the Satilla River; Satilla River, 3 mi. S figured holotype USNM 85153). Lea, 1852, Obs. Unio, 5: 17. Hortense; both Brantly Co. Satilla River, Unio moussonianus Lea Trans. Amer. Camden Co. 1852, Philos. Soc, 10: 268, pi. 18, fig. 22 (Georgia; figured holotype USNM 85168). Lea, 1852, Elliptio (Elliptio) congoraea (Lea) Obs. Unio, 5: 24. Plate 5: 1-8 Unio corvns Lea 1859, Proc. Acad. Nat. Sci.

Phila., 11: 112 ( Buckhead Creek, Burke Co.; Unio Lea 1831, Trans. Amer. Philos. congaraeus Ogeechee River; both Georgia). Lea, 1859, 4: 4 South Soc, 72, pi. 6, fig. (Congaree River, Acad. Nat. Sci. ser. 2, 4: 217, USNM Jour. Phila., Carolina; figured holotype 85693). Lea, pi. 27, fig. 97; figured holotype USNM 84539, 1834, Obs. Unio, 1: 82. from Buckhead Creek. Lea, 1859, Obs. Unio, Unio fulvus Lea 1834, Trans. Amer. Philos. Soc., 7: 35. 5: 96, pi. 13, fig. 39 (South Carolina; figured Unio vestitus Lea 1861, Proc. Acad. Nat. Sci. holotype USNM 85679). Lea, 1834, Obs. Unio, Phila., 13: 393 (Ogeechee River [Georgia]). 1: 208. Lea, 1862, Jour. Acad. Nat. Sci. Phila., ser. 2,

Unio lecontianus Lea 1838, Trans. Amer. Philos. 5: 189, pi. 25, fig. 259; figured holotype USNM Soc., 6: 40, pi. 12, fig. 35 (Canoochee River, 85332. Lea, 1862, Obs. Unio, 9: 11. Liberty Co., Ceorgia; figured holotype USNM Unio dorsatus Lea 1868, Proc. Acad. Nat. Sci. 84852). Lea, 1838, Obs. Unio, 2: 40. Phila., 20: 160 (Catawba River, Nordi Caro- Unio planilateris Conrad 1838, Monography lina). Lea, 1868, Jour. Acad. Nat. Sci. Phila., Atlantic Slope Unionidae • Johnson 309

Width ser. 2, 6: 300, 45, fig. 112; figured holotype Length Height pi. nun mm mm USNM 84494. Lea, 1869, Obs. Unio, 12: 60. 80 Unio strumosus Lea 1872, Proc. Acad. Nat. Sei. Phila., 24: 158 (Yadkin River, North Caro- lina). Lea, 1874, Jour. Acad. Nat. Sci. Phila., ser. 2, 8: 57, pi. 19, fig. 55; figured holotype USNM 85755. Lea, 1874, Obs. Unio, 13: 61.

Description. Shell medium to large, reaching up to 100 mm in length. Outline long, rhomboid, subrhomboid, sometimes almost subtriangular. Valves flat to sub- inflated, thin to solid, inequilateral. An- terior end regularly rounded; posterior end broadly rounded, slightly biangulate or truncate. Ventral margin straight or slightly curved, occasionally a bit arcuate. Dorsal margin short to very short, forming an indistinct angle with the obliquely descend- ing posterior margin. Posterior ridge oc- casionally rather sharp but more often faint to imperceptible, with the slightest hint of a secondary ridge above. Posterior slope rather broad to very broad, with numerous fine ridges that radiate from the upper posterior ridge to the dorsal margin. Umbos rather full, but low, located in the anterior third to fourth of the shell, their sculpture consisting of a few coarse ridges that run parallel to the growth lines. Perios- tracum smooth when young, yellowish with fine to broad green rays, sometimes becoming yellowish green but more often light brown or chestnut, generally smooth on the disk, but roughened toward the margins. Left valve with two heavy, rough pseudocardinal teeth, the more anterior one slightly smaller and a little lower. Hinge line rather short and broad, with two short, slightly curved, granular lateral teeth, the uppermost one sometimes very short and vestigial. Right valve with one chunky, serrated pseudocardinal; one lat- eral tooth. Beak cavities shallow, with a few dorsal muscle scars under the hinge plate. Anterior adductor muscle scars deep, posterior ones and pallial line distinct. Nacre generally white, occasionally light pinkish or purplish. 310 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Range. Southern Atlantic Slope: Ogee- Edisto River System chee River system, Georgia, north to the Edisto River Drainage. South Carolina: Cape Fear River system. North Carolina. Edisto River, Givhans Ferry, 2 mi. NW Specimens Examined Givhans, Dorchester Co.

Ogeechee River System Cooper-Santee River System

Canoochee River Drainage. Georgia: Congaree River Drainage. South Caro- Canoochee River, Bulloch Co. Canoochee lina: Congaree River. River, Liberty Co. (USNM). Canoochee Wateree River Drainage. North Caro- River, E Pembroke (ANSP); Canoochee lina: Catawba River, Tuckasaga Ford [not River, near Clyde (MZUM); both Bryan found] (ANSP). South Carolina: Wateree Co. River, Camden, Kershaw Co. (MZUM). River Drainage. Georgia: Ogeechee Pedee River System Ogeechee River, Shoals, 12 mi. SW War- rington, Warren Co. Ogeechee River, 4 Yadkin River Drainage. South Caro- Co. mi. SW Mitchell, Glascock Ogeechee lina: Yadkin River (ANSP, USNM ). River, 2 mi. S Louisville, Jefferson Co. Cape Fear River System Rocky Creek, 5 mi. SW Waynesboro, Burke Co. Ogeechee River, Scarboro; Ogeechee Cape Fear River Drainage. North Caro- River, bridge, 1 mi. S Dover; Ogeechee lina: Cape Fear River. River, 1.5 mi. SW Oliver; all Screven Co. (all MZUM). Ogeechee River, bridge, 1 Elliptio {Elliptio) dariensis (Lea) mi. E Blitchton; Ogeechee River, Jinks Plate 6: 1-4 Route 16; River, Bridge, Ogeechee Morgan Unio dariensis Lea 1842, Trans. Amer. Philos. 14 mi. Bridge, SE Pembroke; Dolly Lake, Soc, 8: 246, pi. 26, fig. 61 ( [Altamaha River] about 1.5 mi. below Kiterlighter Camp; all near Darien [Mcintosh Co.], Georgia; figured holotvpe USNM 85691). Lea, 1842, Obs. Unio, Co. (a// Bryan MZUM). 3: 84. Savannah River System Unio monroensis Lea 1843, Descriptions Twelve Uniones. (Lake Monroe, Florida). Lea, 1846, Broad River Drainage. Georgia: Broad Trans. Amer. Philos. Soc, 9: 279, pi. 41, fig. Elbert Co. 8; figured holotype USNM 85169. Lea, 1848, River, Huguenot, ( USNM ). Obs. Unio, 4: 37. Savannah River Georgia: Sa- Drainage. Unio wehsterii B. H. Wright 1888, Proc. Acad. Elbert Co. Savannah vannah River, River, Nat. Sci. Phila., p. 113, pi. 2, fig. 2 (Lake Augusta; Spirit Creek, De Bruce; both Rich- Woodruff, Volusia Co., Florida; lectotype USNM selected Occ. mond Co. South Carolina: Savannah River, 125697, by Johnson, 1967, Pap. Moll., 3: 10, pi. 7, fig. 2). 6 mi. W Martin; Savannah River, Johnsons Unio hartwrightii B. H. Wright 1896, Nautilus, 10 mi. Savannah Landing, W Allendale; 9: 121, pi. 2, figs. 4-6 (Lake Beresford [Volusia River, King Jaw Point, 10 mi. WSW Allen- Co.], Florida; holotype USNM 151031, figured Occ. 3: dale (MZUM); all Allendale Co. Georgia: by Johnson, 1967, Pap. Moll., 6, pi. 7, fig. 1 and the type locality further restricted Brier Creek, Chalker Bridge, 6 mi. N on the basis of the original label to: [St. Johns Waynesboro, Burke Co. Savannah River, 7 River], Blue Springs [3 mi. S Lake Beresford, mi. NE Newington, Screven Co. South Volusia Co.], Florida). Carolina: Mouth of Vermezobre Creek (Sa- Description. Shell often large, exceeding vannah River Wildlife Res.) (MZUM); 130 mm in length. Outline subrhomboid Savannah River at Governor Hamilton's or subtrapezoidal. Valves rather flat to 8 mi. S [Mansion, Hardeeville] (ANSP); subinflated, thin but strong, inequilateral. both Jasper Co. Anterior end regularly rounded; posterior Atlantic Slope Unionidak • JoJmson 311

end occasionally a bit produced and Remarks. Elliptic) dariensis (Lea) is slightly biangulate, but more often restricted to the St. Johns River system, obliquely truncated. Ventral margin straight Florida, and the Altamaha River system, or slightly curved. Dorsal margin straight, Georgia, and is close to Elliptic) c. crassi- forming a sharp angle with the obliquely dens (Lamarck), but the shell of dariensis descending posterior margin. Posterior is never as ponderous; the posterior ridge ridge usually very sharp with a faint is consistently much shaqier, especially in secondary ridge above. Posterior slope immature individuals, and the posterior rather broad, but well defined, with nu- slope, though generally wrinkled, does not merous wrinkles on it. Umbos full to in- have the strong radial pattern typical of flated, but rather low, located in the E. c. crassidens. Throughout its range, dariensis is more than E. c. anterior third of the shell, their sculpture compressed downiei even when the consisting of five or six double-looped bars, (Lea) posterior of some individuals of both are not slightly more elevated and angular behind ridge the sinus. Periostracum smooth when very acute. dariensis is also close to E. young, yellowish with fine green rays, be- Elliptic) which is found in the coming darker greenish yellow, or dark congaraea (Lea), to the Fear chestnut or blackish, generally smooth on Ogeechee River, Georgia, Cape but is the disk but sometimes roughened, espe- River, North Carolina, congaraea more variable. It is often more rhomboid cially on the posterior slope. Left valve with two heavy, rough than dariensis. In congaraea the posterior is less and to be pseudocardinal teeth, the more anterior ridge sharp, more prone some of one slightly smaller and a little lower. biangulate, resembling examples Hinge line rather short and broad, with E. crassidens downiei, though the latter are more inflated. Immatures of dariensis two short, slightly curved, granular, lateral tend to have a somewhat chestnut teeth. Right valve with one chunky, ser- perios- tracum with subtle green rays, whereas rated pseudocardinal; one lateral tooth. is often with rather Beak cavities shallow, with a few dorsal congaraea quite yellow bright green rays. muscle scars under the hinge plate. An- In the Altamaha River system, the largest terior adductor muscle scars deep, posterior specimens of E. dariensis are found in the ones and pallial line distinct. Nacre gen- lower Altamaha River. Older specimens erally purple, sometimes white, occasion- have been confused with E. hopetonensis ally yellow. (which replaces E. complanata (Lightfoot) Length Height Width mm mm mm in the main river), but hopetonensis is pro-

130 77 47 House Creek, Bowens Mill, portionally longer and narrower, is without 9 mi. N Ben Fitzgerald, the sharp posterior ridge, and has no Hill Co., Georgia. trace of sculpture on the posterior slope. 98 58 35 As above. Whereas E. hopetonensis is very common 78 55 28 [Altamaha River], near in the Altamaha River, E. dariensis is not. Darien [Mcintosh Co.], Georgia. In a collection made by Clench, Boss and

41 23 14 House Creek, Bowens Mill, Fuller, 10 mi. NE Surrency, Appling Co., 9 mi. N Ben Fitzgerald, Georgia, several hundred hopetonensis Hill Co., Georgia. were collected, but only three dariensis. Anatomy. Discussed by Lea (1863: 404). The especially sharp posterior ridge and Habitat. Lives in sand bars, in fast- relatively thin shell, and the similarity of flowing or even sluggish water. the periostracum, especially as exhibited in 312 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

immatures, suggest that the forms de- figured holotype USNM 84986. Lea, 1863, Obs. 9: 19. scribed from the St. Johns River system Unio, Elliptio mcmichaeli Clench and Turner 1956, Bull. are E. dariensis, even though the St. Johns Florida State Mus., 1: 170, pi. 7, figs. 1-2 and Altamaha river are systems separated ( Choctawhatchee River, 8 mi. W Miller Cross by the St. Marys and Satilla river systems, Roads, Holmes Co., Florida; holotype MCZ which are inhabited by E. c. crassidens and 191922). E. crassidens downiei respectively. Description. Shell medium to large, Peninsular Florida. St. Johns Range. reaching 100 mm in length. Outline sub- River Florida. Southern Atlantic system, elliptical. Valves flat, rather thin but Altamaha River Slope: system, Georgia. solid, inequilateral. Anterior end regularly rounded; posterior end somewhat produced Specimens Examined and biangulate. Ventral margin rather uniformly straight. Dorsal margin straight Altamaha River System or slightly curved, forming an angle with Ocmulgee River Drainage. Georgia: the obliquely descending posterior margin. Rozzy Branch, 4 mi. S Macon; Tobesofkee Posterior ridge fairly well defined near the Creek, 5 mi. S Macon; both Bibb Co. umbos, becoming faint toward the posterior Tucsowhatchee Creek, 5 mi. SW Hawkins- margin, with a second faint ridge above, in a Posterior ville; Cedar Creek, Fountains Mill, 7 mi. ending slight biangulation. SW Hawkinsville; both Pulaski Co. Dick- slope slightly concave and broad, often the with wrinkles sons Creek, 10 mi. NE Fitzgerald, Ben Hill sculptured toward umbos that radiate from the Co. Ocmulgee River, 1.5 mi. S Jackson- posterior ridge to the dorsal Umbos full but located ville; Ocmulgee River, below Lumber City, margin. low, (H. D. Athearn); both Telfair Co. in the anterior fourth of the shell. Perios- tracum smooth the Ohoopee River Drainage. Georgia: and shiny on disk, light Little Ohoopee River, 1 mi. E Kite, Johnson brownish, reddish brown, or yellowish, often with fine Co. Little Ohoopee River, 11 mi. W Swains- green rays when young, with boro; Ohoopee River, 1 mi. E Adrian; becoming uniformly brownish age. Ohoopee River, 0.5 mi. S Norristown; all Left valve with two heavy, rough Emanuel Co. Ohoopee River, above Reids- pseudocardinal teeth, the more anterior one smaller lower. line ville, Tattnall Co. and Hinge very short and rather rather Altamaha River Drainage. Georgia: narrow; two long curved lateral teeth. valve Altamaha River, 10 mi. NE Surrency, slightly Right with Appling Co. Altamaha River, Long Co. one chunky, serrated pseudocardinal; Altamaha River, near Darien, Mcintosh Co. one lateral tooth. Reak cavities very shal- low, with a few dorsal muscle scars under the Anterior adductor muscle Elliptio (Elliptio) fraterna (Lea) hinge plate. scars ones and line Plate 7: 1-5 deep, posterior pallia! distinct. Nacre usually white, sometimes Unio fraternus Lea 1852, Trans. Amer. Philos. pale pink to salmon, or purplish, especially Soc, 10: 263, pi. 16, fig. 15 ([Chattahoochee the River] Columbus [Muscogee Co.], Georgia: toward edges, iridescent posteriorly. USNM 85398; Abbeville District paratype Length Height Width [Savannah River drainage], South Carolina; nun mill mm figured holotype USNM 85396). Lea, Obs. 65 30 20 Abbeville District, South Unio, 5: 19. Carolina. Unio anthonyi Lea 1861, Proc. Acad. Nat. Sei. 60 31 16 As above. Holotype of U. Phila., 13: 41 (Florida). Lea, 1862, Jour. Acad. fraternus Lea.

Nat. Sei. Phila., ser. 2, 5: 197, pi. 27, fig. 266; 50 27 17 As above. Atlantic Slope Unionidae • Johnson 313

Anatomy. Discussed by Lea (1863: 410). Specimens Examined Habitat. Lives in sand liars in large Savannah River System rivers and their smaller tributaries, in swift water. Savannah River Drainage. South Caro- lina: Remarks. Elliptic fraterna (Lea) of the Abbeville District (USNM). Choctawhatchee River system replaces the widely distributed E. c. crassidens (La- Elliptio (Elliptio) waccamawensis (Lea) marck) which is found in both river systems Plate 7: 6, 7 to the Choctawhatchee. It is adjacent Unio waccamawensis Lea 1863, Proc. Acad. Nat. obviously close to crassidens, but the shell Sci. Phila., 15: 193 (Lake Waccamaw, North Acad. Sci. of fraterna is lighter, the outline is sub- Carolina). Lea, 1866, jour. Nat. Phila., ser. 2, 6: 16, pi. 5, fig. 14; figured elliptical, and the shell is consistently more holotype USNM 84437. Ohs. and The Lea, 1867, Unio, elongated compressed. posterior 11: 20. Simpson, 1914, Cat. Naiades, 2:620. ridge is less acute and the posterior slope Description. Shell generally small, not is broader. In the Atlantic drainage, fra- more than 60 oc- terna resembles some forms of E. crassi- reaching mm, though casionally reaching almost 90 mm in length. dens downiei and E. congaraea, but the Outline elongated rhomboid. Valves in- shell of fraterna is consistently thinner and flated or subinflated, thin, inequilateral. more compressed. Anterior end regularly rounded; posterior The discontinuous distribution of popu- end slightly biangulate and pointed toward ations of fraterna was used as a reason for the base. Ventral margin quite straight. establishing mcmichaeli by Clench and Dorsal margin rather short and straight, Turner (1956: In the United States 171). meeting the obliquely descending posterior National Museum are from the specimens margin in a usually distinct angle. Hinge Chattahoochee River, Columbus, Muscogee ligament long and low. Posterior ridge This is near the head- Co., Georgia. locality very high, decidedly angular, with some- River waters of the Choctawhatchee times a faint second ridge above it ending system, and it is likely that fraterna spread behind in a broad point. Posterior slope into the Chattahoochee River in this obliquely truncate, often with wrinkled vicinity, as is suggested here in Chapter 1. sculpture. Umbos neither high nor full, Further, commingling of the waters of the located in the anterior quarter of the shell, upper Chattahoochee and Savannah rivers their sculpture not observed. Periostracum rather with delicate is attested to by a number of species com- smooth, growth lines, sometimes mon to both drainages, besides fraterna. generally smoky-green, yellow- ish or often with dark Clench and Turner claim that mcmichaeli brownish, green rays over the entire surface. is larger than fraterna and that oc- Left valve with two, occasionally three, casional specimens, "show salmon color- stumpy pseudocardinal teeth, one in front ation [nacre]," but a comparison of the of the other, often of almost equal height. and all available series indicate holotypes Hinge line very short and narrow, two that the can not be distinguished specimens rather short straight lateral teeth. Right morphologically. valve with two parallel pscudocardinals, restricted Range. Apalachicolan region: the posterior one inclined to be narrow and River to the Choctawhatchee system, triangular, occasionally broken into two Florida, and the upper Chattahoochee teeth, the more anterior tooth low and River, Georgia, of the Apalachicola River vestigial; one lateral tooth. Reak cavities system. Southern Atlantic Slope: upper very shallow, with a few dorsal muscle Savannah River system, South Carolina. scars. Anterior and posterior adductor 314 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

beside Lake 1 mi. muscle scars and pallial line distinct. Nacre canal, Waccamaw, generally bluish white or pink, sometimes NNW Dupree Landing; all Columbus Co. yellowish, somewhat iridescent. Elliptio {Elliptio) complanata (Lightfoot) Length Height Width mm mm mm Plate 8: 1-6 88 45 28 A 1 mi. drainage canal, Mya complanata (Lightfoot) 1786, Catalogue of NNW Landing, Dupree the Portland Museum, p. 100, no. 2190 (Mary- Columbus Co., North Car- land) refers to: Lister, 1686, Synopsis Method- olina. icae Conchyliorum, pi. 150, fig. 5 (Virginia; Oxford Univ. Mus. 61 34 22 As above. figured specimen [probably lost] teste Dance, 1966, Shell Collecting, p. 54 32 22 Lake Waccamaw, [town 292). Type locality restricted to the Potomac of] Lake Waccamaw, Co- River, Washington, District of Columbia [ap- lumbus Co., North Caro- proximately opposite Fairfax Co., Virginia] by lina. Johnson, 1948, Nautilus, 62: 36. In citing Light- foot rather than Solander (Johnson loc. cit.) as the authority for diis species I have followed Habitat. Lives in sand, in sluggish water. Rehder (1967: 2, 18). Remarks. waccamaicensis (Lea), Elliptio Unio violaccus Spengler 1793, Skrivter af Natur- River which is restricted to the Waccamaw historie Selskabet [Copenhagen], 3: 55 (Norda- system, most closely resembles E. congaraea merika; holotype, Copenhagen Museum; figured (Lea) from the nearby Cape Fear River by Haas, 1913, Vidensk. Meddr. Dansk Naturh. Foren. [Copenhagen], 65: 51, text fig.). system, though waccamaicensis is easily Unio purpureas Say 1817, Nicholson's Encyclo- distinguished from it by being consistently pedia, 2 [no pagination], pi. 3, fig. 1 (Dela- with a more elongated, decidedly more ware and Schuylkill Rivers [presumably near acutely angled posterior ridge. In both Philadelphia, Philadelphia Co., Pennsylvania], ANSP species, the posterior slope has the ten- type, [lost]). Unio rarisulcata Lamarck 1819, Hist. Nat. des dency to have wrinkled sculpture, but the 1 : le lac Animaux sans Vertebres, 6 ( ) 72 ( of waccamaicensis tends to periostracum Champlain [Vermont]; type, Cab. of M. Du- and is more in- be more intensely green fresne [lost]). clined to be rayed. Unio coarctata Lamarck 1819, Hist. Nat. des

sans 6 1 : 73 riviere Specimens of unionids from Lake Wacca- Animaux Vertebres, ( ) (la Hudson [New York]; holotype, Paris Museum, maw have always been greatly corroded figured by Johnson, 1969, Nautilus, 83: 53, fig. and of their eonchologieal characters many 5). but H. D. Athearn has obfuscated, recently Unio purpurascens Lamarck 1819, Hist. Nat. des taken of 1 : les rivieres some more perfect examples Animaux sans Vertebres, 6 ( ) 73 ( of M. waccamaicensis from the obviously hos- de l'etat de New Yorck [sic]; type, Cab. Valenciennes [lost]; type of var. 1) and c, Paris pitable environment of a canal that was Museum, the latter selected as lectotype by to the road on the west side dug parallel Johnson, 1969, Nautilus, 83: 60, fig. 6 du lac of the lake. Champlain [New York]). Range. Southern Atlantic Slope: re- Unio rhombula Lamarck 1819, Hist. Nat. des Animaux sans Vertebres, 6 ( 1 ) : 74 ( Senegal; stricted to the Waccamaw River system, [erroneous] holotype, Geneva Museum, figured North Carolina. by Delessert, 1841, Rec. Coq. de Lamarck, pi. 12, fig. 8). Specimens Examined Unio carinifera Lamarck 1819, Hist. Nat. des

1 : riviere Animaux sans Vertebres, 6 ( ) 74 (la Waccamaw River System Hudson de l'etat de New York; holotype, Paris Museum, figured by Johnson, 1969, Nautilus, Waccamaw River Drainage. Nortlt 83: 52, fig. 4). Carolina: Lake Waccamaw, [town of] Lake Unio georgina Lamarck 1819, Hist. Nat. des 0.5 mi. Waccamaw; Lake Waccamaw, W Animaux sans Vertebres, 6 (1): 74 (le lac [town of] Lake Waccamaw; Drainage George [New York]; holotype, Paris Museum, Atlantic Slope Unionidae • Johnson 315

figured by Johnson, 1969, Nautilus, 83: 53, [//no cuvierianus Lea 1852, Trans. Amer. Philos. fig. 3). Soc, 19: 263, pi. 16, fig. 16 (Washington Co., Unio glabrata Lamarck 1819, Hist. Nat. des Georgia; figured holotype USNM 85401). Lea,

1 : Animaux sans Vertebres, 6 ( ) 75 (la riviere 1852, Obs. Unio, 5: 19. de l'Ohio; [erroneous] holotype, Paris Museum, U/i io errans Lea 1856, Proc. Acad. Nat. Sci. Phila., figured by lolmson, 1969, Nautilus, 83: 53, 8: 262 (Tobesaufke [Tobesofkee] Creek, near fig- 7). Macon [Bibb Co.], Georgia). Lea, 1858, Jour. 17/1/0 sulcidens Lamarck 1819, Hist. Nat. des Acad. Nat. Sci. Phila., ser. 2, 4: 60, pi. 9, fig.

1 : USNM 85541. Animaux sans Vertebres, 6 ( ) 77 ( une riviere 42; figured holotype Lea, 1858, du Connecticut; holotype, Paris Museum, Obs. Unio, 6: 60. Schunglkill [Schuylkill River], Pennsylvania; Unio oicinus Lea 1856, Proc. Acad. Nat. Sci. paratype, Geneva Museum, figured by Deles- Phila., 8: 262 (Swift Creek, near Macon [Bibb sert, 1841, Rec. Coq. de Lamarck, pi. 12, fig. 3. Co.], Georgia). Lea, 1858, Tour. Acad. Nat. (7/1/0 virginiana Lamarck 1819, Hist. Nat. des Sci. Phila., ser. 2, 4: 61, pi. 9, fig. 43; figured USNM 85543. Obs. : holotype Lea, 1858, Unio, Animaux sans Vertebres, 6 ( 1 ) 79 (la riviere 6: 61. Potowmac [Potomac] en Virginie; holotype, Geneva Museum, figured by Delessert, 1841, Unio geminus Lea 1856, Proc. Acad. Nat. Sci. 8: 262 Burke Rec. Coq. de Lamarck, pi. 12, fig. 4. Phila., (Buckhead Creek, Co., Acad. Nat. Sci. l7/i/o aurata Rafinesque 1820, Ann. Gen. Sci. Phys. Georgia). Lea, 1858, Jour. ser. 2, 4: 10, (Bruxelles), 5: 295 (la riviere Hudson [New Phila., 63, pi. fig. 45; figured USNM 84856. Lea, 1858, Obs. Unio, York]; type not in ANSP [lost]). holotype 6: 63. Unio fluviatilis Green 1827, Jour. Maclurian Unio abbevillensis Lea 1857, Proc. Acad. Nat. Lyceum (Philadelphia), 1: 41. [no locality; Sci. Phila., 9: 84 (Abbeville District [Savannah type lost.] River drainage], South Carolina). Lea, 1857, Mya rigida Wood 1828, Index Testaceologicus, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 51, pi. Supplement, pi. 1, fig. 10 (no locality; type 6, fig. 34; figured holotype USNM 85583. Lea, [lost]). Wood, 1856, op. cit., ed. Hanley, p. 1858, Obs. Unio, 6: 51. 200, pi. 1, supp. fig. 10. Unio percoarctatus Lea 1857, Proc. Acad. Nat. Unio griffithianus Lea 1834, Trans. Amer. Philos. Sci. Phila., 9: 85 (Catawba River, Gaston Co., Soc, 5: 103, pi. 15, fig. 46 (South Carolina; North Acad. Nat. USNM Carolina). Lea, 1862, Jour. figured holotype 85610). Lea, 1834, Sci. Phila., ser. 2, 5: 59, pi. 3, fig. 206; figured Obs. Unio, 1: 215. holotype USNM 85529. Lea, 1862, Obs. Unio, l7/i/o complanatus subinflatus Conrad 1835, New 8: 63. Fresh Water Shells United States, Appendix, Unio wheatleyi Lea 1857, Proc. Acad. Nat. Sci.

p. 5, pi. 9, fig. 2 ( Savannah River [Augusta, Phila., 9: 85 (Catawba River, Gaston Co., Richmond Co., Georgia]; syntype ANSP North Carolina). Erroneously thought pre- 119985; and Congaree River, South Carolina). occupied by Lea and changed to: Conrad, 1838, Monographv Unionidae, no. 11, Unio catawbensis Lea 1861, Jour. Acad. Nat. Sci. ser. 5: p. 97, pi. 54, fig. 1. Phila., 2, 54, pi. 1, fig. 200; figured holotype USNM 85547. Lea, 1861, Obs. Unio, Unio jejunus Lea 1838, Trans. Amer. Philos. Soc, 8: 58. 6: 9, pi. 4, fig. 9 (Roanoke River [road between Unio insuhus Lea 1857, Proc. Acad. Nat. Sci. Winton and Tarborough, North Carolina] ) ; Phila., 9: 86 (Roanoke River, Weldon [Halifax [Wateree River], near Camden, [Kershaw Co.], Co.], North Carolina). Lea, 1862, Jour. Acad. South Carolina, figured holotype USNM 85475. Nat. Sci. Phila., ser. 2, 5: 53, pi. 1, fig. 199; Lea, 1838, Obs. Unio, 2: 9. figured holotype USNM 85644. Lea, 1862, Unio roanokensis Lea Trans. Amer. Philos. 1838, Obs. Unio, 8: 57. 6: 21 River Soc, 27, pi. 8, fig. (Roanoke [road Unio spadiceus Lea 1857, Proc. Acad. Nat. Sci. between and Norfolk, Virginia; Tarborough, Phila., 9: 86 (Deep River, Gulf [Chatham Co.]; North Carolina] figured holotype USNM 85423; mountain streams; both North Carolina). Lea, Altamaha [River], Georgia). Lea, 1838, Obs. 1862, Jour. Acad. Nat. Sci. Phila., ser. 2, 5: 55, Unio, 2: 27. pi. 1, fig. 201; figured holotype USNM 85251, Un/'o fuliginosus Lea 1845, Proc. Amer. Philos. from mountain streams, North Carolina. Lea, Soc, 4: 164 (Cobb's Creek, near Philadelphia 1862, Obs. Unio, 8: 59. [Philadelphia Co.], Pennsylvania). Lea, 1848, Unio macer Lea 1857, Proc Acad. Nat. Sci. Trans. Amer. Philos. Soc, 10: 78, pi. 7, fig. 19; Phila., 9: 86 (Roanoke River, Weldon [Halifax figured holotype USNM 85665. Lea, 1848, Co.], North Carolina). Lea, 1862, Jour. Acad. Obs. Unio, 4: 52. Nat. Sci. Phila., ser. 2, 5: 202, pi. 29, fig. 271; 316 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

figured holotype USNM 85427, labeled; Neuse 62, pi. 4, fig. 210; figured holotype USNM River, near Raleigh [Wake Co.], North Caro- 85002. Lea, 1860, Obs. Unio, 8: 66. lina. Lea, 1863, Obs. Unio, 9: 24. Unio postellii Lea 1858, Proc. Acad. Nat. Sci. Unio contractus Lea 1857, Proc. Acad. Nat. Sci. Phila., 10: 165 (Randall's Creek, near Colum- Phila., 9: 86 (Roanoke River, Weldon [Halifax bus [Muscogee Co.]; Carter's Creek, Baldwin Co.], North Carolina). Lea, 1862, Jour. x\cad. Co.; both Georgia). Lea, 1859, Jour. Acad. Nat. Sei. Phila., ser. 2, 5: 203, pi. 29, fig. 272: Nat. Sci. Phila., ser. 2, 4: 214, pi. 26, fig. 94; figured holotype USNM 86102. Lea, 1862, figured holotype USNM 85470 from Carter's Obs. Unio, 9: 25. Creek. Lea, 1859, Obs. Unio, 7: 32. Unio virens Lea 1857, Proc. Acad. Nat. Sci. Unio roswellensis Lea 1858, Proc. Acad. Nat. Sci. Phila., 9: 169 (Georgia). Lea, 1858, Jour. Phila., 10: 165 (Chattahoochee River, Roswell, Acad. Nat. Sci. Phila., ser. 2, 4: 80, pi. 16, Cobb Co., Georgia). Lea, 1859, Jour. Acad. fig. 60; figured holotype USNM 85544. Lea, Nat. Sci. Phila., ser. 2, 4: 205, pi. 24, fig. 87; 1858, Obs. Unio, 6: 80. figured holotype USNM 85467. Lea, 1859, Unio savannahensis Lea 1857, Proc. Acad. Nat. Obs. Unio, 7: 23. Sci. Phila., 9: 169 (Savannah River, Rrantley's Unio hurkensis Lea 1859, Proc. Acad. Nat. Sci. Mill [not located], Washington Co., both Phila., 11: 112 (Buckhead Creek, Burke Co., Georgia; Santee Canal, South Carolina; Sugar Georgia). Lea, 1859, Jour. Acad. Nat. Sci. Creek, Mecklenburg Co., North Carolina). Lea, Phila., ser. 2, 4: 215, pi. 27, fig. 95; figured 1858, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: holotype USNM 85998. Lea, 1859, Obs. Unio, 80, pi. 16, fig. 61; figured holotype USNM 7: 33. 85404 from Savannah River. Lea, 1858, Obs. Unio hallenbeckii Lea 1859, Proc. Acad. Nat. Unio, 6: 81. Sci. Phila., 11: 170 (Flat Rock Creek; Four Unio suhflavus Lea 1857, Proc. Acad. Nat. Sci. Mile Creek; both near Columbus [Muscogee Phila., 9: 170 (Walnut Creek above Macon Co.], Georgia). Lea, 1860, Jour. Acad. Nat. [Bibb Co.], Georgia); Lea, 1858, Jour. Acad. Sci. Phila., ser. 2, 4: 328, pi. 51, fig. 154; Nat. Sci. Phila., ser. 2, 4: 90, pi. 19, fig. 70; figured holotype USNM 85537, from Black Dirt figured holotype USNM 85562. Lea, 1858, Creek, near Columbus [Muscogee Co.], Georgia. Obs. Unio, 6: 90. Lea, 1860, Obs. Unio, 8: 10. Unio fumatus Lea 1857, Proc. Acad. Nat. Sci. Unio haldwinensis Lea 1859, Proc. Acad. Nat. Phila., 9: 171 (Chattahoochee River, near Sci. Phila., 11: 170 (Carter's Creek, Baldwin Columbus [Muscogee Co.], Georgia; Hospaliga Co., Georgia). Lea, 1860, Jour. Acad. Nat. Creek, Alabama, [and] Georgia). Lea, 1858, Sci. Phila., ser. 2, 4: 330, pi. 51, fig. 155; Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 88, pi. figured holotype USNM 85420. Lea, 1860, 18, fig. 68; figured holotype USNM 85552, Obs. Unio, 8: 12. from Chattahoochee River. Lea, 1858, Obs. Unio salcbrosus Lea 1859, Proc. Acad. Nat. Sci. Unio, 6: 88. Phila., 11: 170 (Flat Rock Creek; Bull Creek; Unio subniger Lea 1857, Proc. Acad. Nat. Sci. Chattahoochee River; all near Columbus [Mus- Phila., 9: 172 (Flint River, near Macon cogee Co.], Georgia). Lea, 1860, Jour. Acad. [County], Georgia). Lea, 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 332, pi. 52, fig. 157; Nat. Sci. Phila., ser. 2, 4: 196, pi. 22, fig. 79; figured holotype USNM 85574, from Flat Rock figured holotype USNM 85560. Lea, 1859, Creek. Lea, 1860, Obs. Unio, 8: 14. Obs. Unio, 7: 14. Unio raeensis Lea 1859, Proc. Acad. Nat. Sci. Unio neusensis Lea 1858, Proc. Acad. Nat. Sci. Phila., 11: 171 (Chattahoochee [River], near Phila., 10: 41 (Neuse River, 6 miles from [East] Columbus [Muscogee Co.] and Rae's Creek

Raleigh [Wake Co.], North Carolina). Lea, [not located] , Georgia). Lea, 1869, Jour. 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 5: Acad. Nat. Sci. Phila., ser. 2, 4: 331, pi. 52, fig. 60, pi. 4, fig. 208; figured holotype USNM 156; figured holotype USNM 85571. Lea, 1860, 85465. Lea, 1860, Obs. Unio, 8: 64. Obs. Unio, 8: 13. Unio purus Lea 1858, Proc. Acad. Nat. Sci. Phila., Unio latus Lea 1859, Proc. Acad. Nat. Sci. Phila., K>: 41 (Neuse River, 6 miles from [East] 11: 171 (Savannah River, near Savannah Raleigh [Wake Co.], North Carolina). Lea, [Chatham Co.], Georgia). Lea, 1860, Jour. Sci. 1859, Jour. Acad. Nat. Phila., ser. 2, 5: Acad. Nat. Sci. Phila., ser. 2, 4: 334, pi. 53, 61, pi. 4, fig. 209: figured holotype USNM fig. 159; figured holotype USNM 85898. Lea, 85003. Lea, 1860, Obs. Unio, 8: 65. 1860, Obs. Unio, 8: 16. Unio exactus Lea 1858, Proc. Acad. Nat. Sci. Unio (iiiadratus Lea 1859, Proc. Acad. Nat. Sci. Phila., 10: 41 (Neuse River, 6 miles from [East] Phila., 11: 172 (Carter's Creek, [Baldwin Co.]; Raleigh [Wake Co.], North Carolina). Lea, Factory Creek, near Columbus [Muscogee Co.]; 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 5: both Georgia). Lea, 1860, Jour. Acad. Nat. Atlantic Slope Untonidae • Johnson 317

Sci. Phila., ser. 2, I 338, pi. 54, fig. 163; 19; figured holotype USNM 85672 from Rocky figured holotype USNM 85718 from Factor) Run. Lea, 1867, Obs. Unio, 11: 25. Creek. Lea, 1860, Obs. Unio, 8: 20. Unio gastonensis Lea 1863, Proc. Acad. Nat. Sci. Unio squamous Lea 1861, Proc. Acad. Nat. Sci. Phila., 15: 191 (Mine Creek, Gaston Co., North Phila., 13: 391 (North Carolina). Lea, 1862, Carolina). Lea, 1866, Jour. Acad. Nat. Sci. lour. Acad. Nat. Sci. Phila., ser. 2, 5: 200, pi. Phila., ser. 2, 6: 20, pi. 6, fig. 18; figured holo- 28, fig. 269; figured holotype USNM 85530. type USNM 85409. Lea, 1867, Obs. Unio, Lea, 1863, Obs. Unio, 9: 22. 11: 24. Unio rostrum Lea 1861, Proc. Acad. Nat. Sci. Unio quadrilatcrus Lea 1863, Proc. Acad. Nat. Phila., 13: 391 (Davidson Co., North Caro- Sci. Phila., 15: 192 (Abbeville District [Savan- lina). Lea, 1862, Jour. Acad. Nat. Sci. Phila., nah River drainage], South Carolina; Neuse ser. 2, 5: 201, pi. 29, fig. 270; figured holotype River, near [6 miles East of] Raleigh [Wake USNM 85556. Lea, 1863, Obs. Unio, 9: 23. Co.]; Catawba River; both North Carolina). Unio northamptonensis Lea 1861, Proc. Acad. Lea, 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, Nat. Sci. Phila., 13: 392 (Connecticut River, 6: 5, pi. 1, fig. 1; figured holotype USNM Northampton [Hampshire Co.]; [Connecticut 85385 from the Neuse River. Lea, 1867, Obs. River] Springfield [Hampden Co.]; both Massa- Unio, 11: 9. chusetts. [Connecticut River] below Hartford Unio indefinilus Lea 1863, Proc. Acad. Nat. Sci. [Hartford Co.], Connecticut; Neuse River, Phila., 15: 192 (Long Creek, Mecklenburg Co.; North Carolina). Lea, 1862, Jour. Acad. Nat. Neuse River, near [6 miles East of] Raleigh Sci. Phila., ser. 2, 5: 190, pi. 25, fig. 260; [Wake Co.]; both North Carolina). Changed figured holotype USNM 85527 from the Con- to: necticut River, Northampton [Hampshire Co.], Unio indefinitus Lea 1866, Jour. Acad. Nat. Sci. Massachusetts. Lea, 1863, Obs. Unio, 9: 12. Phila., ser. 2, 6: 15, pi." 4, fig. 12; figured Unio decumbens Lea 1861, Proc. Acad. Nat. Sci. holotype USNM 85388, from Long Creek. Lea, Phila., 14: 40 (Alabama). Lea, 1862, Tour. 1867, Obs. Unio, 11: 19. Acad. Nat. Sci. Phila., ser. 2, 5: 87, pi. 12, fig. Unio cistclliformis Lea 1863, Proc. Acad. Nat. 236; figured holotype USNM 86150 [not seen]. Sci. Phila., 15: 192 (Neuse River, near [6 Lea, 1862, Obs. Unio, 8: 91. Is E. complanata, miles East of] Raleigh [Wake Co.], North Caro- fide Frierson, 1922, Nautilus, 36: 44. lina). Lea, 1866, Jour. Acad. Nat. Sci. Phila., Unio rcdeighensis Lea 1863, Proc. Acad. Nat. Sci. ser. 2, 6: 19, pi. 6, fig. 17; figured holotype Phila., 15: 191 (Neuse River, 6 mi. East of USNM 85533. Lea, 1867, Obs. Unio, 11: 23. Raleigh [Wake Co.], North Carolina). Lea, Unio mediocris Lea 1863, Proc. Acad. Nat. Sci. 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: 6, Phila., 15: 192 (Neuse River, [6 miles East of] pi. 1, fig. 2; figured holotype USNM 85592. Raleigh [Wake Co.], North Carolina). Lea, Lea, 1867, Obs. Unio, 11: 10. 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: Unio aberrans Lea 1863, Proc. Acad. Nat. Sci. 24, pi. 7, fig. 22; figured holotype USNM Phila., 15: 191 (Neuse River, 6 mi. East of 85611. Lea, 1866, Obs. Unio, 11: 28. Raleigh [Wake Co.], North Carolina). Lea, Unio perlucens Lea 1863, Proc. Acad. Nat. Sci. 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: 7, Phila., 15: 193 (Six Runs, Sampson Co., North pi. 1, fig. 3, figured holotype USNM 85600. Carolina). Lea, 1866, Jour. Acad. Nat. Sci. Lea, 1867, Obs. Unio, 11: 1. Phila., ser. 2, 6: 18, pi. 5, fig. 16; figured Unio weldonensis Lea 1863, Proc. Acad. Nat. Sci. holotype USNM 85076. Lea, 1867, Obs. Unio, Phila., 15: 191 (Roanoke River, Weldon [Hali- 11: 22. fax Co.], North Carolina). Lea, 1866, Jour. Unio curatus Lea 1863, Proc. Acad. Nat. Sci. Acad. Nat. Sci. Phila., ser. 2, 6: 11, pi. 3, fig. Phila., 15: 193 (Sugar Creek [Mecklenburg 8; figured holotype USNM 85407. Lea, 1867, Co.], North Carolina). Lea, 1866, Jour. Acad. Obs. Unio, 11: 15. Nat. Sci. Phila., ser. 2, 6: 23, pi. 7, fig. 21; Unio mecklenbergensi.s Lea 1863, Proc. Acad. type, C. M. Wheatley collection in ANSP; the Nat. Sci. Phila., 15: 191 (near Charlotte, only specimen located, ANSP 127200, is not Mecklenberg [sic] Co., North Carolina). Lea, figured type. Lea, 1867, Obs. Unio, 11: 27. 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: 17, Unio protensus Lea 1865, Proc. Acad. Nat. Sci. pi. 5, fig. 15, figured holotype USNM 85419. Phila., 17: 33 (North Carolina). Lea, 1868, Lea, 1867, Obs. Unio, 11: 21. Tour. Acad. Nat. Sci. Phila., ser. 2, 6: 256, pi. Unio chathamensis Lea 1863, Proc. Acad. Nat. 31, fig. 71; figured holotype USNM 85994, Sci. Phila., 15: 191 (Rocky Run, Chatham Co., from Eno River, near Hillsboro, Orange Co., North Carolina; James River, near Richmond North Carolina. Lea, 1868, Obs. Unio, 12: 16. [Henrico Co.], Virginia). Lea, 1866, Jour. Unio lazarus Sowerby 1868 in Reeve, Conch. Acad. Nat. Sci. Phila., ser. 2, 6: 21, pi. 6, fig. Iconica 16, Unio, pi. 68, fig. 348 (Abbeville 318 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

District [Savannah River drainage], South Caro- Co.; Long Creek, Gaston Co.; both North Caro- lina) non Lea, 1852. lina). Lea, 1874, Jour. Acad. Nat. Sci. Phila., Unio beaverensis Lea 1868, Proc. Acad. Nat. Sci. ser. 2, 8: 49, pi. 17, fig. 47; figured holotype Phila., 20: 161 (Beaver Creek [Gaston Co.]; USNM 85590, from Irwin's Creek. Lea, 1874, Long Creek [Gaston Co.]; both North Caro- Obs. Unio, 13: 53. lina. Carter's Creek [Baldwin Co.], Georgia). Unio differtus Lea 1872, Proc. Acad. Nat. Sci. Lea, 1868, Jour. Acad. Nat. Sci. Phila., ser. 2, Phila., 24: 158 ([Savannah River], Georgia ?). 6: 297, pi. 44, fig. 109; figured holotype USNM Lea, 1874, Jour. Acad. Nat. Sci. Phila., ser. 2, 85598 from Beaver Creek. Lea, 1869, Obs. 8: 42, pi. 14, fig. 39; figured holotype USNM Unio, 12: 57. 85399. Lea, 1874, Obs. Unio, 13: 46. Unio nubilus Lea 1868, Proc. Acad. Nat. Sci. Unio sirfjparallclus Lea 1872, Proc. Acad. Nat. Phila., 20: 161 (Paw Creek, Mecklenburg Co., Sci. Phila., 24: 158 (Irwin's Creek [Mecklen- North Carolina). Lea, 1868, Jour. Acad. Nat. Sci. burg Co.]; Fox River [not located]; both North Phila., ser. 2, 6: 298, pi. 44, fig. 110; figured Carolina). Lea, 1874, Jour. Acad. Nat. Sci. holotvpe USNM 85417. Lea, 1869, Obs. Unio, Phila., ser. 2, 8: 46, pi. 16, fig. 44; figured 12: 58. holotype USNM 85418, localities not separated. Unio datus Lea 1868, Proc. Acad. Nat. Sci. Phila., Lea, 1874, Obs. Unio, 13: 50. 20: 161 (Paw Creek [Mecklenburg Co.]; Beaver Unio oblongus Lea 1872, Proc. Acad. Nat. Sci. Co. [ = Creek, Gaston Co.]; Long Creek Phila., 24: 158 (Irwin's Creek, Mecklenburg [Gaston Co.]; all North Carolina). Lea, 1868, Co., North Carolina). Lea, 1874, Jour. Acad. Jour. Acad. Nat. Sci. Phila., ser. 2, 6: 299, pi. Nat. Sci. Phila., ser. 2, 8: 52, pi. 18, fig. 50; 44, fig. Ill; figured holotype USNM 85097 figured holotype USNM 85572. Lea, 1874, labeled, "Paw Creek, Beaver Co., North Caro- Obs. Unio, 13: 56. lina." Lea, 1869, Obs. Unio, 12: 59. Unio curvatus Lea 1872, Proc. Acad. Nat. Sci. Unio humerosus Lea 1868, Proc. Acad. Nat. Sci. Phila., 24: 159 (Pfeiffers Pond [Charlotte], Phila., 20: 161 (Charlotte, Mecklenburg Co., Mecklenburg Co., North Carolina). Lea, 1874, North Carolina). Lea, 1868, Jour. Acad. Nat. Jour. Acad. Nat. Sci. Phila., ser. 2, 8: 47, pi. Sci. Phila., ser. 2, 6: 301, pi. 45, fig. 113; 13, fig. 35; figured holotype USNM 85550. figured holotype USNM 85414. Lea, 1869, Lea, 1874, Obs. Unio, 13: 42. Obs. Unio, 12: 61. Unio invinensis Lea 1872, Proc. Acad. Nat. Sci. Unio uhareensis Lea 1868, Proc. Acad. Nat. Sci. Phila., 24: 159 (Irwin's Creek, [Mecklenburg Phila., 20: 145 (Uharee [Uwharree] River, Co.], North Carolina). Lea, 1874, Tour. Acad. Montgomery Co., North Carolina). Lea, 1868, Nat. Sci. Phila., ser. 2, 8: 44, pi. 15, fig. 42; Jour. Acad. Nat. Sci. Phila., ser. 2, 6: 304, pi. figured holotype USNM 85680. Lea, 1874, 46, fig. 116; figured holotype USNM 85576. Obs. Unio, 13: 48. Lea, 1869, Obs. Unio, 12: 63. Unio subsquamosus Lea 1872, Proc. Acad. Nat. Unio tortuosus Sowerby 1868, Conch. Iconica, 16, Sci. Phila., 24: 160 (Yadkin River, North Unio, pi. 65, fig. 330 (Maryland; figured holo- Carolina). Lea, 1874, Jour. Acad. Nat. Sci. type BMNH 74.12.11.25). Phila., ser. 2, 8: 47, pi. 16, fig. 45; figured Unio santeensis Lea 1871, Proc. Acad. Nat. Sci. holotype USNM 85557. Lea, 1874, Obs. Unio, Phila., 23: 193 (Santee Canal, South Carolina; 13: 51. Oconee River, Georgia). Lea, 1874, Tour. Acad. Unio infuscus Lea 1872, Proc. Acad. Nat. Sci. Nat. Sci. Phila., ser. 2, 8: 20, pi. 6, fig. 17; Phila., 24: 160 (Fox River [not located]; figured holotype USNM 85635, from Santee Irwin's Creek, Mecklenburg Co.; both North Canal. Lea, 1874, Obs. Unio, 13: 24. Carolina). Lea, 1874, Jour. Acad. Nat. Sci. Unio yadkinensis Lea 1872, Proc. Acad. Nat. Sci. Phila., ser. 2, 8: 51, pi. 17, fig. 49; figured Phila., 24: 156 (Yadkin River, near Salisbury holotype USNM 85587 from Fox River. Lea, [Rowan Co.], North Carolina). Lea, 1874, 1874, Obs. Unio, 13: 55. Jour. Acad. Nat. Sci. Phila., ser. 2, 8: 32, pi. Unio ratus Lea 1872, Proc. Acad. Nat. Sci. Phila., 10, fig. 29; figured holotype USNM 85387. 24: 160 (Neuse River, North Carolina). Lea, Lea, 1874, Obs. Unio, 13: 36. 1874, Tour. Acad. Nat. Sci. Phila., ser. 2, 8: 52, Unio ampins Lea 1872, Proc. Acad. Nat. Sci. pi. 18, fig. 51; figured holotype USNM 85382. Phila., 24: 157 (Irwin's Creek, Mecklenburg Lea, 1874, Obs. Unio, 13: 56. Co., North Carolina). Lea, 1874, Jour. Acad. Unio basalis Lea 1872, Proc. Acad. Nat. Sci. Nat. Sci. Phila., ser. 2, 8: 39, pi. 13, fig. 36; Phila., 24: 161 (Carter's Creek, near Colum- figured holotype USNM 85591. Lea, 1874, bus [Muscogee Co.], Georgia). Lea, 1874, Obs. Unio, 13: 43. Jour. Acad. Nat. Sci. Phila., ser. 2, 8: 48, pi. Unio ligatus Lea 1872, Proc. Acad. Nat. Sci. 16, fig. 46; figured holotype USNM 85703. Phila., 24: 157 (Irwin's Creek, Mecklenburg Lea, 1874, Obs. Unio, 13: 52. Atlantic Slope Umomdae • Johnson 319

Unio dissimilis Lea 1872, Proc. Acad. Nat. Sci. Pilsbry, 1891, Nautilus, 4: 127. Pilsbry, 1892, Phila., 24: 161 (Long Creek, Gaston Co.; Proc. Acad. Nat. Sci. Phila., 44: 132, pi. 7, Pfeiffers Pond, [Charlotte], Mecklenburg Co.; figs. 4-6; figured holotype ANSP 63094. both North Carolina). Lea, 1874, Jour. Acad. Unio palliatus 'RaveneV Simpson 1900, Proc. Nat. Sci. Phila., ser. 2, 8: 53, pi. 18, fig. 52; United States Natl. Mus., 22: 730. [nomen figured holotype USNM 86019 from Long nudum]. Listed under the synonymy of Unio Creek. Lea, 1874, Obs. Unio, 13: 57. errans Lea. Unio cirratus Lea 1874, Proc. Acad. Nat. Sci. Unio pullatus majusculus De Gregorio 1914, II Phila. for 1873, 25: 422 (Abbeville District Naturalista Siciliano, 22: 57, pi. 8, figs, a-d [Savannah River drainage], South Carolina). (Connecticut; type Palermo Museum, Sicily ser. Lea, 1874, Jour. Acad. Nat. Sci. Phila., 2, [not seen] ). 8: 55, pi. 19, fig. 53; figured holotype USNM Unio complanatus mainensis Rich 1915, Science, 85584. Obs. 13: 59. Lea, 1874, Unio, n. s., 42: 580 ( Songo Pond about 3 miles Unio subolivaceus Lea 1874, Proc. Acad. Nat. South of Rethel [Oxford Co.], Maine; type, Sci. Phila. for 1873, 25: 422 (Catawba River; [location not known]). Fox River [not located]; Yadkin River; all Elliptio violaceus (Spengler). Ortmann, 1919, Acad. Nat. North Carolina). Lea, 1874, Jour. Mem. Carnegie Mus., 8: 94, pi. 8, figs. 4, 5. Sci. ser. 8: 56; Phila., 2, 57, pi. 20, fig. figured Elliptio strigosus (Lea), partim. Clench and 85564. Obs. holotype USNM Lea, 1874, Unio, Turner, 1956, Bull. Florida State Mus., 1: 165. 13: 61. Elliptio complanatus (Solander). Clarke and Unio Lea Proc. Acad. Nat. Sci. infulgens 1874, Berg, 1959, Cornell Univ. Exp. Sta. Mem. 367, Phila. for 25: 422 (Stewart's Pond, Union 1873, p. 21, figs. 26-28. Athearn and Clarke, 1962, North Acad. Co., Carolina). Lea, 1874, Jour. Natl. Mus. Canada, Bull., 183, p. 22, pi. 1, Nat. Sci. Phila., ser. 2, 8: 60, pi. 21, fig. 59; figs. 5, 6; pi. 4, figs. 7, 8. figured holotype USNM 85403. Lea, 1874, Obs. Unio, 13: 64. Description. Shell large, often reaching Unio corneus Lea 1874, Proc. Acad. Nat. Sci. over 100 mm in length. Outline generally Phila. for 1873, 25: 423 ([Chattahoochee long, trajDezoidal, sometimes decidedly River] Columbus [Muscogee Co.]; Marietta rhomboid, occasionally subelliptical. Valves [Cobb Co.]; both Georgia. Abbeville District oc- [Savannah River drainage], South Carolina). generally flat or subinflated, though Lea, 1874, Jour. Acad. Nat. Sci. Phila., ser. 2, casionally considerably inflated, thin to 8: USNM 59, pi. 20, fig. 58; figured holotype solid. Anterior end regularly rounded; 85580. Lea, 1874, Obs. Unio, 13: 63. posterior end usually broader and some- Unio dooleijcnsis Lea 1874, Proc. Acad. Nat. Sci. what biangulate. Ventral margin straight Phila. for 1873, 25: 424 (Flint River, Dooley or rjarallel to the Co., Georgia; Abbeville District [Savannah slightly arcuate, roughly River drainage] South Carolina). Lea, 1874, long straight or slightly curved dorsal mar- Jour. Acad. Nat. Sci. Phila., ser. 2, 8: 64, pi. gin, which forms an obtuse angle with the USNM 85538 22, fig. 60; figured holotype obliquely descending joosterior margin. from Flint River. Lea, 1874, Obs. Unio, 13: 68. Hinge ligament prominent. Posterior ridge Unio gesnerii Lea 1874, Proc. Acad. Nat. Sci. usually broad, double and rounded, some- Phila. for 1873, 25: 424 (Uchee River [Russell times rather in a biangu- Co., Alabama], near Columbus [Muscogee angular, ending base. Posterior broad Co.], Georgia). Lea, 1874, Jour. Acad. Nat. lation near the slope unin- Sci. Phila., ser. 2, 8: 65, pi. 22, fig. 64; figured and unsculptured. Umbos low and Obs. holotype USNM 85670. Lea, 1874, Unio, flated, their sculpture consisting of strong 13: 69. ridges that run nearly parallel to the Unio invenustus Lea 1874, Proc. Acad. Nat. Sci. growth lines and are carried back to the Phila. for 1873, 25: 424 ( [Chattahoochee River] nucleus behind as delicate radial lirae. Columbus [Muscogee Co.]; Russell Co.; both often Georgia. Irwin's Creek [Mecklenburg Co.], Surface with irregular growth lines, North Carolina). Lea, 1874, Jour. Acad. Nat. nearly smooth in young shells, becoming Sci. Phila., ser. 2, 8: 66, pi. 22, fig. 62; figured rougher in old ones. Periostracum brown- from holotype USNM 85704 Columbus, Georgia. to ish, or yellowish green, greenish brown, Lea, 1874, Obs. Unio, 13: 70. almost black, often with green rays over Unio (Arconaia) provancheriana Pilsbry 1890, entire surface. Nat. Canadienne, 20: 171 (locality unknown). the 320 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Left valve with two stumpy pseudo- most widely distributed and abundant cardinal teeth, one in front of the other, unionid in the Atlantic Slope region, ex- the more anterior one triangular, the tending from the upper Altamaha River, hinder one generally not much elevated Georgia, to northern Canada. It is a very above the hinge line. Hinge line short and variable species, but most nineteenth cen- narrow; two long, straight, granular lateral tury authors agreed on uniting all of the teeth. Right valve with one chunky, ser- ecophenotypes north of about the latitude rated pseudocardinal, with a vestigial tooth of Washington under complanata. South

in front of it; one lateral tooth. Beak of Washington, Isaac Lea, especially when cavities very shallow, with a few dorsal in his seventies and eighties, applied spe- muscle scars. Anterior and posterior adduc- cific names to the ecophenotypes of com- tor muscle scars and pallial line all distinct. planata and icterina as assiduously as Nacre generally purplish, though often Bourguignat and Locard did to the com- white, or slightly orange; iridescent, espe- mon European unionids, with the same cially toward the margins. confusion resulting. Simpson (1914, 2: to the 651 ) did a great deal rectify Length Height Width mm mm mm synonymy of complanata. This species, al- 154 82 44 Tar River, Bruce, 9 mi. though variable in outline, degree of in- NW Greenville, Pitt Co., flation, and color of both periostracum and North Carolina. nacre, is generally trapezoidal in outline, 99 50 28 Savannah River, 7 mi. NE with the valves rather and Newington, Screven Co., compressed, Georgia. when swollen, the greatest width is in the 77 43 20 Turkey Creek, 4 mi. NE region of the posterior ridge. In large Allentown, Wilkinson Co., rivers such as the Savannah, Neuse, Tar, Georgia. Roanoke, lower Potomac, and Connecti- Anatomy. See Ortmann (1912a: 269). cut, specimens reach maximum size, and Discussed and figured by Reardon (1929: tend to be subelliptical in outline with 11, pi. 4, figs. 1-10). flattened valves, whereas in smaller rivers Breeding season. Ortmann (1919: 104) and tributaries they tend to be more found gravid females from April 26 through rhomboid in outline and more inflated. July 16 during several years in Pennsyl- Walter (1956: 266, 270) regarded the vania; the breeding season was restudied small creek form as E. c. complanata and by Matteson (1948b: 704), who found that called the large river form E. complanata in the Ocqueoc River, lower Michigan, it roanokcnsi.s (Lea), even though both extended from the middle of June to the forms were found together at a number of middle of July. the stations. Habitat. Found in lakes, ponds, small In the Altamaha River system E. com- streams, and large rivers on nearly every planata can be confused with E. hope- type of substrate, though it appears to tonensis (Lea) (under which see: Remarks prefer sand. Throughout its range it is on p. 325) which replaces it in the Alta- sometimes the only unionid found at some maha River proper. It can also be confused stations, and at others it is generally found with E. icterina (Conrad) (under which in greater numbers than any other species see: Remarks on p. 328) with whose save where its distribution overlaps Ellip- several ecophenotypes it is often found, tio icterina (Conrad). south of the White Oak River, North Caro- Remarks. Elliptio complanata (Light- lina. In Georgia, complanata can not be foot), which is found at scattered localities confused with Uniomerus tetralasmus in the Apalachicola River system, is the (Lea) with its acutely angled meeting of Atlantic Slope Unionidae • Johnson 321

the dorsal and posterior margins and its Ogeechee River System satiny periostracum. These characteristics Ogeechee River Drainage. Georgia: I become somewhat less constant in the Ogeechee River, Shoals, Warren Co. Carolinas and northward, and complanata Ogeechee Creek, 4 mi. SW Mitchell, Glas- can be confused with it. However, tetra- cock Co. Big Creek, 3 mi. SE Louisville; lasmus tends to be more quadrate and to Williamson Swamp Creek, Bartow; Nails exhibit more swelling of the valves, and is Creek, 2 mi. S Bartow; Rocky Creek, 2 mi. always unrayed. S Wadley; Big Creek, 3 mi. E Louisville; The specimens of E. complanata from all Jefferson Co. Mill Creek, 5 mi. E Mid- Lake Creek, at Stone Mountain, De Kalb ville; Buckhead Creek, 14 mi. W Waynes- Co., Georgia, are among just a few records boro; Buckhead Creek, 10 mi. SW of Unionidae collected above the Fall Line Waynes- boro; all Burke Co. Ogeechee River, in Georgia. Unfortunately, I misidentified Scarboro (MZUM); Ogeechee River, them as hopetonensis for Raulerson and bridge, 1 mi. S Dover (MZUM); both Burbank (1962, 9: 39). Screven Co. Ogeechee River, bridge, 1 mi. Range. Apalachicolan region: restricted E Blitchton (MZUM); Ogeechee River, to the Apalachicola River system. Atlantic Jinks Bridge, Route 16 (MZUM); Ogeechee Slope region: Altamaha River system, River, Morgan Bridge, 14 mi. SE Pembroke Georgia, north to the St. Lawrence River (MZUM); all Bryan Co. system, Canada. Interior Basin: westward to Lake Superior, also parts of the Hudson Savannah River System Bay drainage. Discussed in detail and Broad River Drainage. Georgia: Cadya illustrated by Matteson (1948a: 13, fig. 2). Creek [not located] (MZUM). Broad Specimens Examined River, 1.5 mi. S Bell; Broad River, Hugue- not; both Elbert Co. Altamaha River System Savannah River Drainage. South Caro- lina: Abbeville Ocmulgee River Drainage. Georgia: Co. (USNM). Turkey Lake Creek at Stone Mountain, De Kalb Creek, 7 mi. NW Edgefield, Edgefield Co. Walnut Creek, E Macon; Tobesofkee Co. Georgia: Savannah River, Augusta, Creek, 5 mi. S Macon (USNM); both Bibb Richmond Co. (MZUM). South Carolina: Lower Three 8 mi. Barn- Co. Flat Creek, 2 mi. S Perry; Big Creek, Runs Creek, SW 3 mi. S Henderson; both Houston Co. Buck well, Barnwell Co.; Savannah River, 6 mi. Creek, 5 mi. NW Hawkinsville; Limestone W. Martin; Savannah River, Johnsons Creek, 4.3 mi. E Hawkinsville; Cedar Landing, 10 mi. W Allendale; Savannah Creek, 5 mi. WSW Hawkinsville; Tusca- River, King Jaw Point, 10 mi. WSW Allen- whatchee Creek, 5 mi. SW Hawkinsville; dale (MZUM); all Allendale Co. Georgia: all Pulaski Co. Bluff Creek, 8 mi. NW Brier Creek, Keysville (MZUM); Mill mi. Burke Abbeville, Wilcox Co. Dicksons Creek, Race, 2 N Sardis; both Co. mi. 10 mi. NW Fitzgerald, Ben Hill Co. Savannah River, 7 NE Newington, Oconee River Drainage. Georgia: Car- Screven Co. South Carolina: Mouth of ters Creek, Baldwin Co. Turkey Creek, Vermezobre Creek (Savannah River Wild-

4 mi. life Res. Co. . Savannah NE Allentown, Wilkinson Co. Ford ) , Jasper ( MZUM ) Branch, 4 mi. W Dublin; Turkey Creek, River, Savannah, Chatham Co. 9 mi. W Dublin; both Laurens Co. COMBAHEE Rr\T:R SYSTEM Ohoopee River Drainage. Georgia: Ohoopee River, 4 mi. S Wrightsville, John- Salkehatchee River Drainage. South son Co. Carolina: Lemon Creek, 2.5 mi. S Bam- 322 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

berg; Little Salkehatchee River, 4 mi. N Black River System both Co. Ehrhardt; Bamberg Whippy Black River Drainage. South Carolina: 2.5 mi. NE Crocketville, Swamp Creek, Sammy Swamp Creek, 6 mi. S Paxville, Hampton Co. Clarendon Co.

Edisto River System Pedee River System River South Caro- Edisto River Drainage. Soutli Carolina: Lynches Drainage. lina: 2 mi. NE Snake Swamp Creek, Orangeburg Co., 6 Lynches River, Bishopville, Lee Co. mi. NE Bamberg, Bamberg Co. N. Fork Yadkin River North Caro- Edisto River, Orangeburg, Orangeburg Co. Drainage. lina: Yadkin Yadkin Co. Edisto River, Givhans Ferry, 2 mi. NW River, Boomville, Co. Givhans, Dorchester Co. Yadkin River, Salisbury, Rowan (MZUM). Ditch, Buffalo Creek, 6 mi. E Ashley River System Concord, Cabarrus Co. (MZUM). Uwhar- rie River, Montgomery Co. (USNM). Ashley River Drainage. South Carolina: Stewarts Pond, Union Co. (USNM). [Ashley River] near Summersville, Dor- chester Co. Waccamaw River System Waccamaw River North Cooper-Santee River System Drainage. Carolina: Lake Waccamaw, [town of] Lake Saluda River Drainage. South Caro- Waccamaw, Columbus Co. Soutli Caro- lina: Saluda River, 2.3 mi. above Ware lina: Ditch, Waccamaw River, 1.25 mi. Shoals, Abbeville Co. SE Conway, Horry Co. Waccamaw River, Broad River Drainage. Tennessee: Wachasaw Landing, 2 mi. W Murrells Green River, 5 mi. ENE Mill Spring, Polk Inlet, Georgetown Co. Co. South Carolina: Broad Headwaters, Cape Fear Rwer System River, SE Reidsville, Spartenburg Co. River North Carolina: Congaree River Drainage. South Caro- Deep Drainage. Sandv Creek, Co. lina: Congaree River, Columbia, Richland Randolph (MZUM). Gulf Co. Rocky Run (USNM); Deep River, (Lea); both Chatham Co. Catawba River Drainage. North Caro- Cape Fear River Drainage. North Caro- lina: Catawba River, Bridgewater, Burke lina: Haw River, 1.25 mi. NE Benaja, Co. Leppers Creek, Lincolnton, Lincoln Co. Buffalo Creek, Co. Mine Creek; Beaver Creek; both Gas- Rockingham (MZUM). 1 mi. E Greensboro, Guilford Co. ton Co. Paw Creek; Stewarts Creek; Irwins (MZUM). Morgan Creek, 1 mi. SE Creek; Sugar Creek; Ashleys Creek; Long Hill, Co. New Creek; Catawba River; Bissels Pond, Char- Chapel Orange (MZUM). Hope River, Burke Forest, Durham Co. lotte; Pfeiffers Pond, Charlotte; Elias Pond, River, 11 mi. N Sanford, 10 mi. from Charlotte; all Mecklenburg Co. (MZUM). Rocky Chatham Co. Fear River, Carlos; Soutli Carolina: Branch, Little Dutchmans Cape Fear River, Kinnon; both Cumber- Creek, Rockhill, York Co. Cape land Co. (both MZUM). Ashe River, Wateree River Drainage. South Caro- Pender Co. Greenfield Mill Pond, Wil- lina: Wateree River, 2.5 mi. W Camden, New Hanover Co. Kershaw Co. mington, (USNM). South River Drainage. North Carolina: Santee River Drainage. South Caro- Six Runs, Sampson Co. (USNM). lina: Poplar Creek, Schulers Fish Pond, near Santee State Park, Orangeburg Co., Neuse River System Wilsons Landing, 5 mi. NW Pineville; Neuse River Drainage. North Carolina: Santee Canal; both Berkeley Co. North Flat River, 5.5 mi. S Roxboro Center, Atlantic Slope Unionidae • Johnson 323

1 Person Co. (SI) Eno River, 1.75 mi. ESE Roanoke River, Ironto (USNM); both Ilillshoro Center S Little ( 117); Fork, River, Montgomery Co. Black Water River, 0.5 mi. 104 Lake 12 mi. Franklin N Schley ( ); Michie, Co. (MZUM). North Carolina: NNE Durham Center (75); Neuse River, 6 Roanoke River, Washington Co. mi. ENE Durham Center all Dur- (64); Chowan River System ham Co. Neuse River, 10.5 mi. NE Raleigh (49); Walnut Creek, 4.75 mi. WSW Meherrin River Drainage. Virginia: Raleigh (32); Swift Creek, 3 mi. WSW Meherrin River, 2 mi. SW Grandy, Bruns- Garner (21); all Wake Co. Little River, wick Co. Three Creek, near Emporia, 1.25 mi. NW Ragley (2); Neuse River, Greensville Co. (MZUM). North Caro- 3.5 mi. NNE Smithfield (26); both John- lina: Meherrin River, Murfreesboro, Hert- ston Co. Neuse River, Cliffs of Neuse State ford Co. (USNM). Park, 2 mi. NW Seven Springs, Wayne Co. Nottoway River Drainage. Virginia: Neuse River, 13.5 mi. WSW Kingston Cen- Tommeheton Lake, near Camp Pickett, ter, Lenoir Co. Neuse River, Streets Ferry, Nottoway Co. Waqua Creek, 2 mi. SE 8.5 mi. NNW New Bern, Craven Co. Rawlings; Nottoway River, 3 mi. E Rawl- Trent River, near Pollocksville, Jones Co. ings; both Brunswick Co. (MZUM). Blaekwater River Drainage. Virginia: Swift Creek, Rolling Mills, Chesterfield Co. Pamlico River System Nansemond River, Nansemond Co. Tar River Drainage. Nortli Carolina: Chowan River Drainage. Virginia: Tar River, Providence, Granville Co. Sandy Chowan River, Edenhouse, 13 mi. E Wind- Creek, Franklin Co. Stony Creek, Nash- sor, Bertie Co. ville, Nash Co. [Beach Swamp Creek] James River System Enfield, Halifax Co. Swift Creek, Whit- River Calf- akers ( MZUM ) ; Tar River, Tarboro; Tar James Drainage. Virginia: River, Old 3.5 mi. W all pasture River (Conrad, 1846: 407). North Sparta, Pinetops; = Edgecomb Co. Tar River, Bruce, 9 mi. [ Maury] River, Lexington, Rockbridge Co. 2 mi. NW Greenville; Mill Run, Greenville; both (USNM). Rivanna River, W Pitt Co. Columbia, Fluvanna Co. James River, opposite Maidens, Goochland Co. James Roanoke River System River, near Cartersville, Cumberland Co. Dan River Drainage. Virginia: Aarons James River, Richmond, Henrico Co. Creek, 3 mi. W Buffalo Lithia Springs; Dan (MZUM). River; both Halifax Co. Bluestone Creek, Chiekahominy River Drainage. Vir- near Clarksville, Mecklenburg Co. ginia: Chickahominy River, 4 mi. S Ash- Roanoke River Drainage. Virginia: land, Hanover Co. Roanoke 4 mi. SW N. Fork River, Elliston; York River System

1 Walter (1956) marked on a map of the North Anna River Drainage. Virginia: Upper Neuse River some of the 136 stations he Mine Run; Church River; both Orange Co., made his in 1950-51. He collected during survey {both MZUM). E. complanata at forty of them. A selection of these with the station numbers included are Rappahannock River System presented here, since the localities have never been published. According to the author, most Rapidan River Drainage. Virginia: of the collections were made with a 1/8 inch Mountain Run, Orange Co. (USNM). mesh which accounts for the scraper net, paucity Rappahannoek River Drainage. Vir- of specimens in most of the lots and also why ginia: River, 1 mi. S Rem- this and other species were not found at more Rappahannock of the stations. ington, Faquier Co. Mountain Run, 3 mi. N 324 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Lignum, Culpeper Co. Rappahannock wedge-shaped or slightly biangulate. Ven- River, Fredericksburg, Spotsylvania Co. tral margin generally straight or slightly arcuate. Dorsal margin long and straight, Occoquan Creek System tending to be a bit winglike where it joins Occoquan Creek Drainage. Virginia: the obliquely descending posterior margin Rull Run, 3 mi. N Catharpin; Broad Run, in either a sharp or imperceptible angle. [Fairfax Co.], 3 mi. W Manassas; Kettle Hinge ligament prominent. Posterior ridge Prince Run, 1.5 mi. N Nokesville; all usually rather broad, single, and narrowly William Co. rounded above, becoming double below and in a wide at or Potomac River System ending biangulation above the posterior base. Posterior slope Potomac River West Vir- Drainage. rather broad and unsculptured. Umbos River, 1 mi. below Inter- ginia: Cacapon low, slightly inflated, located in the an- Co. Potomac mont, Hampshire (USNM). terior third of the shell, their sculpture River, Brosius, Co. Canal at Harp- Morgan consisting of several bars. Disk rather flat, ers Ferry, Co. (MZUM). Virginia: Jefferson or slightly concave when a slight umbonal- [North River of S Fork Shenandoah ventral sulcus is present. Periostracum River], Weyers Cave, Augusta Co. Pass- brownish or yellowish green to olive green, age Creek, 2 mi. SW Seven Fountains often with green rays in young individuals, (USNM); N Fork Shenandoah River, E becoming rough and brownish or blackish Woodstock; both Shenandoah Co. Mary- in matures. land: Potomac River, 0.75 mi. W Point of Left valve with two stumpy pseudo- Rocks, Frederick Co. Virginia: Potomac cardinal teeth, one in front of the other, River, near Great Falls; Little Hunting the more anterior one the hinder Creek, near Mt. Vernon (MZUM); both triangular, the Fairfax Co. one generally not much elevated above hinge line. Hinge line short and narrow, two lateral teeth. Elliptio {Elliptio) hopetonensis (Lea) long, straight, granular valve with one serrated Plate 9: 1-2 Right chunky, pseudocardinal, with a vestigial tooth in Unio hopetonensis Lea 1838, Trans. Amer. Philos. front of it; one lateral tooth. Beak cavities Soc, 6: 29, pi. 9, fig. 24 ( [Altamaha River] Hopeton, near Darien [Mcintosh Co.], Georgia; very shallow, with a few dorsal muscle figured holotype USNM 85391). Lea, 1838, scars. Anterior and posterior adductor Obs. Unio, 2: 29. Simpson, 1914, Cat. Naiades, muscle scars and pallia] line all distinct. 2: 668. Nacre white or pinkish, occasion- Unio inusitatis Lea 1859, Proc. Acad. Nat. Sci. generally and iridescent, Phila., 11: 171 (Swift Creek, below Macon ally purplish especially [Bibb Co.], Georgia). Lea, 1860, Jour. Acad. toward the margins. Nat. Sci. Phila., ser. 2, 4: 333, pi. 52, fig. 158; Length Height Width figured holotype USNM 85531. Lea, 1860, mm mm mm Obs. 8: 15. Cat. Unio, Simpson, 1914, Naiades, 150 75 38 Altamaha River, 3 mi. NW 2: 670. Everett City, Glynn Co., Georgia. Description. Shell large, often exceed- 103 55 37 Ohoopee River, 3 mi. N ing 150 mm in length. Outline subrhom- Leman, Emanuel Co., boid, narrower anteriorly. Valves rather Georgia. flat to subinflated, occasionally quite in- 84 43 20 Altamaha River, 4 mi. NE Geor- flated, the greatest inflation being about Jesup, Wayne Co., gia. midway along the posterior ridge, they are thin to solid, inequilateral. Anterior end Anatomy. Discussed by Lea (1863: 415). regularly rounded; posterior end broader, Habitat. Lives in sand or sandy mud in Atla\ii< Slope Unionidai'. • Johnson 325

the Altamaha River and its major tribu- Co. Oconee River, 2.5 mi. N Glenwood, taries. Wheeler Co.

Remarks. Elliptic) hopetonensis (Lea), Ohoopee River Drainage. Georgia: which is restricted to the Altamaha River Ohoopee River, Norristown; Ohoopee system, is sympatric with Elliptic com- River, 3 mi. N Leman; both Emanuel Co. phinata (Lightfoot), which replaces hope- Ohoopee River, above Reidsville, Tattnall tonensis in the smaller streams of the Co. (ANSP). system. While E. complanata is very vari- Altamaha River Drainage. Georgia: able in both outline and degree of inflation, Altamaha River, 7 mi. N Ha/.lehurst, Jeff E. hopetonensis is quite constant in outline Davis Co. Altamaha River, 10 mi. N and in its lack of inflation. When some- Baxley, Appling Co. Altamaha River, what inflated, as it is at the limits of its "Riverside Park," 4 mi. N Jesup, Wayne range, the greatest degree is near the mid Co. Altamaha River, 3 mi. NE Everett portion of the posterior ridge. The posterior City, Glynn Co. Altamaha River, near Fort ridge is better defined in hopetonensis and Barrington; Altamaha River, Hopeton, near the long dorsal margin tends to render most Darien; both Mcintosh Co. individuals wedge-shaped. The large river form of E. complanata is Elliptio (Elliptio) icterina (Conrad) admittedly very close to hopetonensis, but Plate 9: 3-10 in river svstems other than the Altamaha, Plate 10: 1-3 where the flat-sided form is found, large Undo icterinus Conrad, [May] 1834, New Fresh are more typical intergrades generally Water Shells United States, p. 41, pi. 6, fig. present. 5 (muddy shore, Savannah River, opposite Augusta [Richmond Co.], Georgia; figured E. hopetonensis (Lea) has been confused holotype ANSP 41381). Published in May, with older of E. dariensis specimens (Lea), teste Conrad, 1853, Proc. Acad. Nat. Sci. Phila., but the latter has a proportionally higher 6: 244, and not disputed by Lea, 1854, Proc. Acad. Nat. Sci. 7: 336-349. Conrad, shell, with a sharper posterior ridge, often Phila., 1836, Monography Unionidae, no. 4, p. 39, pi. with sculpture on the posterior slope. 18, fig. 2. Southern Atlantic re- Range. Slope: Unio raveneli Conrad, [May] 1834, New Fresh stricted to the lower Altamaha River Water Shells United States, p. 39, pi. 6, fig. small system, Georgia. 4 (Wateree Canal; since found in the creeks near Cooper River; vicinity of Santee all South Carolina. Two ANSP Specimens Examined Canal; syntypes 41370; the smaller one agrees with Conrad's but is not the Altamaha River System description, figured specimen, which appears to be Elliptio complanata ( Light- The second is lanceo- Ocmulgee River Drainage. Georgia: foot). specimen Elliptio lata non Unio ravenelianus Lea, 1834). Swift Creek below Macon, Bibb Co. (Lea), Unio ivatereensis Lea 1836, Synopsis Unionidae, Hawkinsville; Cedar Ocmulgee River, p. 31. New name for Unio raveneli Conrad, Creek, Fountains Mill, 7 mi. SW Hawkins- 1834, non Unio ravenelianus Lea, 1834. As out Proc. United ville; both Pulaski Co. House Creek, pointed by Simpson, 1900, States Natl. Mus., 22: 748, this change was Bowens Mill, 10 mi. N Fitzgerald, Ben Hill unnecessary. Co. River, 1.5 mi. S Ocmulgee Jackson- Unio confertus Lea, [August or September] 1834, ville, Telfair Co. Trans. Amer. Philos. Soc, 5: 103, pi. 16, fig. Little Ocmulgee River Drainage. Geor- 47 (Santee Canal, South Carolina; type not in USNM [presumed lost]). Lea, 1834, Obs. gia: Gum Swamp Creek, 1 mi. N McRae, Unio, 1: 215. Published in August or Septem- Telfair Co. ber 1834, teste Lea, 1854, Proc. Acad. Nat. Oco- Oconee River Drainage. Georgia: Sci. Phila., 7: 244. nee River, 8 mi. SW Soperton. Treutlen Unio lugubris Lea 1834, Trans. Amer. Philos. 326 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Soc, 6: 30, pi. 9, fig. 25 ([Altamaha River], Phila., ser. 2, 5: 59, pi. 3, fig. 207; figured Hopeton, near Darien [Mcintosh Co.], Georgia; holotype USNM 85077 from the Catawba figured holotype USNM 85638). Lea, 1838, River. Lea, 1862, Obs. Unio, 8: 63. Obs. Unio, 2: 30, non Say, 1832. Unio obnubilus Lea 1857, Proc. Acad. Nat. Sci. Unio geddingsianus Lea 1840, Proc. Amer. Philos. Phila., 9: 169 (Buckhead Creek, Burke Co., Soc., 1: 285 (Congaree River, South Carolina). Georgia). Lea, 1858, Jour. Acad. Nat. Sci. Lea, 1842, Trans. Amer. Philos. Soc, 8: 202, Phila., ser. 2, 4: 84, pi. 17, fig. 64; figured pi. 11, fig. 15; figured holotype USNM 85650. holotype USNM 85646. Lea, 1858, Obs. Unio, Lea, 1842, Obs. Unio, 3: 40. 6: 84. Unio fuscatus Lea 1843, Desc. Twelve Uniones Unio opacus Lea 1857, Proc Acad. Nat. Sci. (Black Creek, Florida). Lea, 1846, Trans. Phila., 9: 169 (Buckhead Creek, Burke Co., Amer. Philos. Soc, 9: 277, pi. 40, fig. 4; figured Georgia). Lea, 1858, Jour. Acad. Nat. Sci. holotvpe USNM 85243. Lea, 1848, Obs. Unio, Phila., ser. 2, 4: 86, pi. 18, fig. 66; figured 4: 35. holotype USNM 85546. Lea, 1858, Obs. Unio, Unio occulta* Lea 1843, Desc. Twelve Uniones 6: 86. (Black Creek; Lake Monroe; both Florida). Unio similis Lea 1857, Proc. Acad. Nat. Sci. Lea, 1846, Trans. Amer. Philos. Soc, 9: 279, Phila., 9: 169 (Buckhead Creek, Burke Co., pi. 41, fig. 7; figured holotype USNM 85247, Georgia). Lea, 1858, Jour. Acad. Nat. Sci. from Black Creek. Lea, 1848, Obs. Unio, 4: 37. Phila., ser. 2, 4: 91, pi. 19, fig. 71; figured Unio limatulus Conrad 1849, Proc. Acad. Nat. holotype USNM 85653. Lea, 1858, Obs. Unio, Sci. Phila., 4: 154 (Savannah River, [Georgia]; 6: 91. type not in ANSP, [presumed lost]). Conrad, Unio sublatus Lea 1857, Proc. Acad. Nat. Sci. 1850, Jour. Acad. Nat. Sci. Phila., ser. 2, 1: 276, Phila., 9: 169 ([Chattahoochee River], Uchee pi. 37, fig. 9. Conrad, 1853, Proc. Acad. Nat. Bar, below Columbus [Muscogee Co.], Georgia). Sci. Phila., 6: 251. Lea, 1858, Jour. Acad. Nat. Sci. Phila., ser. 2, Unio tuomeyi Lea 1852, Trans. Amer. Philos. 4: 82, pi. 16, fig. 62; figured holotype USNM Soc, 10: 256, pi. 13, fig. 4 (Abbeville District 85897. Lea, 1858, Obs. Unio, 6: 82. [Savannah River drainage], South Carolina; Unio viridicatus Lea 1857, Proc. Acad. Nat. Sci. figured holotype USNM 85669). Lea, 1852, Phila., 9: 170 (Buckhead Creek, Burke Co., Obs. Unio, 5: 12. Georgia). Lea, 1858, Jour. Acad. Nat. Sci.

Unio whiteianus Lea 1852, Trans. Amer. Philos. Phila., ser. 2, 4: 87, pi. 18, fig. 67; figured Soc, 10: 258, pi. 14, fig. 8 (near Savannah holotype USNM 85551. Lea, 1858, Obs. Unio, [Chatham Co.], Georgia; figured holotype 6: 87. USNM 85658). Lea, 1852, Obs. Unio, 5: 14. Unio tetricus Lea 1857, Proc. Acad. Nat. Sci. Unio barrattii Lea 1852, Trans. Amer. Philos. Soc, Phila., 9: 170 (Flint River, near Albany 10: 256, pi. 13, fig. 5 (Abbeville District [Dougherty Co.], Georgia). Lea, 1859, Jour. [Savannah River drainage], South Carolina; Acad. Nat. Sci. Phila., ser. 2, 4: 195, pi. 22, figured holotype USNM 86010). Lea, 1852, fig. 78; figured holotype USNM 85655. Lea, Obs. Unio, 5: 12. 1859, Obs. Unio, 7: 13. Unio pullatis Lea 1856, Proc. Acad. Nat. Sci. Unio aequatus Lea 1857, Proc. Acad. Nat. Sci. Phila., 8: 262 (creeks near Columbus [Mus- Phila., 9: 170 (Buckhead Creek, Burke Co., cogee Co.], Georgia). Changed to: Georgia). Lea, 1858, Jour. Acad. Nat. Sci. Unio puUatus Lea 1858, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 89, pi. 19, fig. 69; figured Phila., ser. 2, 4:57, pi. 8, fig. 39; figured holo- holotype USNM 85561. Lea, 1858, Obs. Unio, type USNM 86020. Lea, 1858, Obs. Unio, 6: 89. 6: 57. Unio aquilus Lea 1857, Proc Acad. Nat. Sci. Unio coruscus Could 1856, Proc Boston Soc. Nat. Phila., 9: 172 (Flint River, Macon [County], Hist., 6: 15 (River Saint John's, near Lake Georgia). Lea, 1858, Jour. Acad. Nat. Sci.

Beresford, Florida; measured holotype MCZ Phila., ser. 2, 4: 92, pi. 20, fig. 72; figured 169097, figured by Frierson, 1911, Nautilus, 25, holotype USNM 85993. Lea, 1858, Obs. Unio, pi. 1, figs. 1-3, and by Johnson, 1964, United 6: 92. States Natl. Mus., Bull. no. 239, p. 60, pi. Unio viridiradiatus Lea 1859, Proc. Acad. Nat. 32, fig. 3). Sci. Phila., 11: 154 (Big Uchee River [Creek, Unio micans Lea 1857, Proc. Acad. Nat. Sci. Russell Co., Alabama] near Columbus, Georgia). 9: Phila., 85 (Catawba River, Gaston Co.; Lea, 1860, Jour. Acad. Nat. Sci. Phila., ser. 2, Gulf Deep River, [Chatham Co.]; both North 4: 336, pi. 53, fig. 161; figured holotype USNM Carolina). Lea, 1862, Jour. Acad. Nat. Sci. 86018). Lea, 1860, Obs. Unio, 8: 18. Atlantic Slope Unionidae • Johnson 327

Unto hepaticus Lea 1859, Proc. Acad. Nat. Sci. Phila., 24: 159 (Buckhead Creek, Burke Co.,

Abbeville I River Phila., 11: 151 ( Salkehatchie River, South Ceorgia; District Savannah Carolina). Lea, 1860, Jour. Acad. Nat. Sci. drainage], South Carolina). I. '-a, 1874, Jour. Sci. ser. Phila., ser. 2, 4: 348, pi. 57, fig. 173; figured Acad. Nat. Phila., 2, 8: 43, pi. 14, fig. holotype USNM 85559. Lea, 1860, Obs. Unio, 50; figured holotype USNM 86014, from Buck- 8: 30. head Creek. Lea, 1874, Obs. Unio, 13: 47. Unio viridans Lea 1859, Proc. Acad. Nat. Sci. Unto hastatus Lea 1873, Proc. Acad. Nat. Sci. Phila., 11: 170 (near Columbus [Muskogee Phila., 25: 423 (New Market, Abbeville District Co.], Georgia). Lea, 1860, lour. Acad. Nat. Sci. [Savannah River drainage], South Carolina). Phila., ser. 2, 4: 337, pi. 54, fig. 162; figured Lea, 1874, Jour. Acad. Nat. Sci. Phila., ser. 2, holotype USNM 85579. Lea, 1860, Obs. Unio, 8: 56, pi. 19, fig. 54; figured holotype USNM 8: 19. 86013. Lea, 1874, Obs. Unio, 13: 60. Unio verutus Lea 1859, Proc. Acad. Nat. Sci. Unio fryanus B. H. Wright 1888, Proc. Acad. Phila., 11: 171 (Flat Rock Creek, near Co- Nat. Sci. Phila., p. 113, pi. 2, fig. 1 (Lake lumbus [Muscogee Co.], Ceorgia). Lea, 1860, Ashby, Volusia Co., Florida; figured holotype Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 335, USNM 151032, refigured by Johnson, 1967, pi. 53, fig. 160; figured holotype USNM 85899. Occ. Papers on Moll., 3: 6, pi. 8, fig. 5). Lea, 1860, Obs. Unio, 8: 17. Unto nolani B. H. Wright 1888, Proc. Acad. Nat. Proc. Acad. Nat. Unio ocmulgeensis Lea 1861, Sci. Phila., p. 116, pi. 4, fig. 11 (a creek flow- Sci. Phila., 13: 38 (Little Ocmulgee River, ing into St. Johns River, near Palatka [Putnam Lumber City [Telfair Co.], Georgia). Lea, Co.], Florida; holotype USNM 151030, re- 1862, Jour. Acad. Nat. Sci. Phila., ser. 2, 5: figured by Johnson, 1967, Occ. Papers on Moll., 95, pi. 14, fig. 243; figured holotype USNM 3: 7, pi. 10, fig. 4). 85901. Obs. 8: 99. Lea, 1862, Unio, Unio simpsoni B. H. Wright 1888, Proc. Acad. Unio merceri Lea Proc. Acad. Nat. Sci. 1862, Nat. Sci. Phila., p. 117, pi. 5, fig. 1 (Lake 169 River drain- Phila., 14: (Lee Co., [Flint Woodruff, Volusia Co., Florida; holotype USNM Acad. Nat. age], Georgia). Lea, 1862, Jour. 151038, refigured by Johnson, 1967, Occ. Sci. Phila., ser. 2, 5: 209, pi. 31, fig. 278, Papers on Moll., 3: 8, pi. 8, fig. 2). figured holotype USNM 86057. Lea, 1863, Unio hurtchianus S. H. Wright 1897, Nautilus, Obs. Unio, 9: 31. 10: 137 (St. Marys River, Nassau Co., Florida; Unio lucidus Lea 1863, Proc. Acad. Nat. Sci. lectotype USNM 149653, selected by Johnson, Phila., 15: 192 (Livingston's Creek, Bruns- 1967, Occ. Papers on Moll., 3: 5, pi. 8, fig. 4, wick Co., [NE corner of Columbus Co., Cape possibly the specimen figured by Simpson, 1900, Fear River drainage], North Carolina). Lea, Proc. Acad. Nat. Sci. Phila., p. 80, pi. 4, fig. 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: 9, 8). pi. 2, fig. 6; figured holotype USNM 85242. Unio diazensis S. H. Wright 1897, Nautilus, 11: Lea, 1867, Obs. Unio, 11: 13. 5 (Lake Diaz, Volusia Co., Florida; lectotype Unio livingstonensis Lea 1863, Proc. Acad. Nat. USNM 149652, selected by Johnson, 1967, Occ. Sci. Phila., 15: 192 (Livingston's Creek, Bruns- on Moll., 3: 6, pi. 8, fig. 6). wick Co., [NE corner of Columbus Co., Cape Papers Unio B. H. 1899, Nautilus, 13: Fear River drainage] North Carolina). Lea, dispalans Wright 50 (Suwannee River, Florida; holotype USNM 1866, Jour. Acad. Nat. Sci. Phila, ser. 2, 6: 159986, figured by Simpson, 1900, Proc. Acad. 14, pi. 4, fig. 11; figured holotype USNM Nat. Sci. 1, fig. 9, refigured 85536. Lea, 1867, Obs. Unio, 11: 18. Phila., p. 80, pi. Occ. on Moll., 3: 6, Unto ablatus [sic] Lea 1863, Proc. Acad. Nat. by Johnson, 1967, Papers Sci. Phila., 15: 193 (Long Creek, Gaston Co., pi. 8, fig. 3). Unto B. H. 13: North Carolina). Changed to: singuUris Wright 1899, Nautilus, branch of the Flint Unio oblatus Lea 1866, Jour. Acad. Nat. Sci. 75 (Spring Creek, [a River], Decatur measured holotype Phila., ser. 2, 6: 13, pi. 4, fig. 10; figured Co., Georgia; 1967, Occ. holotype USNM 86001. Lea, 1867, Obs. Unio, USNM 159988, figured by Johnson, 11: 17. Papers on Moll., 3: 8, pi. 5, fig. 7). List Unio radiolus Lea 1871, Proc. Acad. Nat. Sci. EUiptio cylindraccus Frierson 1927, Check North American 29, new name for Phila., 23: 192 (Ogeechee River, Liberty Co., Naiades, p. Unio non 1832. Georgia). Lea, 1874, Jour. Acad. Nat. Sci. lugubris Lea, 1838, Say, B. H. Nautilus, Phila., ser. 2, 8: 21, pi. 6, fig. 18; figured EUiptio maywebbae Wright 1934, 5-8 Seminole holotype USNM 85621. Lea, 1874, Obs. Unio, 48: 28; ibid. 47, pi. 13, figs. (near 13: 25. Springs [3.4 miles NE Sorrento], 15 miles SE Eustis Unto cuspitatus Lea 1872, Proc. Acad. Nat. Sci. of [Lake Co.], Florida, refigured by 328 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Length Height Width Johnson, 1967, Occ. Papers on Moll., 3: 7, mm mm mm pi 10, fig. 3). 98 49 29 Napiers Creek, 6 mi. N Elliptio strigosus (Lea), partim. Clench and Tur- Melntyre, Wilkinson Co., ner, 1956, Bull. Florida State Mus., 1: 165. Georgia. Description. Shell generally small to 81 42 28 Mill Race, 2 mi. N Sardis, Burke medium, seldom reaching over 100 mm Co., Georgia. 88 50 30 Canoochee River, 2 mi. N in length. Outline variable, subquadrate Claxton, Evans Co., Geor- to sometimes rather pointed. subelliptical, gia. Valves subinflated, subsolid to very solid, Habitat. Found in lakes, ponds, small inequilateral. Anterior end regularly streams, and large rivers, in nearly every rounded; posterior end generally biangu- type of substrate. E. icterina (Conrad) is late near the base, though sometimes sometimes found with E. complanata rather produced and pointed. Ventral mar- (Lightfoot) and other Unionidae at a gin straight or slightly arcuate, sometimes given station, but like complanata, it is obliquely descending. Dorsal margin often found alone. In any case, one or the slightly curved or almost straight, meeting other of these species is generally more the descending posterior margin obliquely abundant than any other unionid at a in a more or less distinct angle. Hinge given station. 7 ligament long and low . Posterior ridge Remarks. Elliptio icterina (Conrad), rounded, double. broadly generally faintly which ranges from the Escambia River Posterior flat to concave, slope slightly system in the Apalachicolan region, east sometimes with faint radial very sculpture through northern Florida, and north to the broad and full but present. Umbos very White Oak River of North Carolina, is a the anterior of the low, located in quarter very variable species and a number of several shell, their sculpture consisting of populations have been named, some of double-looped ridges. Disk surface gen- them several times over. While some popu- erally flat, or slightly concave when a slight lations are more or less identifiable, there umbonal-ventral sulcus is present. Perios- is usually a gradual transition of them be- tracum generally fine and shiny, though tween one river system and the next, such sometimes heavy and rough, black, brown- that while specimens from extremes of the bear little an- ish black, or yellowish brown or bright range resemblance to one there to be no at yellow, chestnut, often with numerous very other, appears point which can be established. There fine green rays. subspecies is often Left valve with two stumpy pseudo- a great deal of ecophenotypic vari- ation, even at what to be a cardinal teeth, one in front of the other, appears single station, the extremes usually connected by often of about equal height. Hinge line intergrades. short and narrow; two long straight lateral In the Apalachicolan region, E. icterina teeth. Right valve with two roughly paral- can be confused with E. complanata lel pseudocardinals, the posterior one apt (Lightfoot) and arctata (Conrad). The to be serrated and chunky, the more an- latter is a rare species outside of the Ala- terior one low and vestigial; one lateral bama River system. It is distinctly and con- tooth. Beak cavities shallow, with a very sistently arcuate with compressed valves, few dorsal muscle scars. Anterior and pos- whereas icterina has a generally straight terior adductor muscle scars and pallia! or curved ventral margin, is bluntly or line all distinct. Nacre generally purplish, acutely pointed posteriorly, and when oc- though sometimes salmon, bluish white, or casionally produced post-basally, the valves pinkish, posteriorly iridescent. are somewhat inflated. Elliptio icterina Atlantic Slope Unionedae • Johnson 329

occurs with E. complanata in the Apalach- medial line, the shells of which tend to be icola River system, and it can be dis- heavier, more inflated, and produced post- tinguished from complanata by its less basally. This shell form occurs again in rhomboid, more elongate, often pointed abundance in Buckhead Creek of the shape. Ogeechee River system; Brier Creek, of The most common shape E. icterina the Savannah River system; both Georgia; takes in the Apalaehicolan region is sub- and the Salkehatchie River of South Car- rhomboidal to subelliptical, sometimes olina; all on the Atlantic Slope, which, appearing quite pointed posteriorly if the similar to the Floridian rivers mentioned, triangulated posterior ridge ends near the are known to be rich in carbonates, Text- medial line. The tendency to be pointed figure 4. That the shape of the shell is is more prevalent in specimens from the apparently environmentally controlled is Apalaehicolan region than in those from also illustrated by the close resemblance of the Southern Atlantic Slope. The similarity shells from Magnesia Springs, 3 miles W of the populations of this species from the Hawthorn, Alachua Co., Florida; the head- Chattahoochee River and the upper Savan- waters of the Ogeechee River; and Cedar nah River, first noted in the localities of Spring, 2 mi. SE Bamberg, Bamberg Co., some of the taxa described by Isaac Lea, South Carolina. Though from widely further confirms the commingling of the separated drainage systems, shells from headwaters of these two systems. The shell these stations bear a closer resemblance to form just described includes most of the one another than they do to specimens taxa grouped by Simpson (1914, 2: 661) from other stations in their respective under U. Lea and Clench and The shell form de- tuomei/i" by drainage systems. just Turner (1956: 165-169) under Elliptic scribed includes most of the taxa grouped strigosus (Lea). by Simpson (1914, 2: 641) under Unio Noteworthy are the populations in Moc- obnubilis Lea and some of those (1914, casin Creek of Econfina Creek, Bay Co., 2: 639) under Unio confertus Lea. Florida, on the Gulf Coast: Black Creek, Especially on the Southern Atlantic Florida, St. Marys and Canoochee rivers, Slope, often in the lower parts of rivers, Georgia, all on the Atlantic Slope; they specimens of E. icterina tend to be less resemble one another more than they do subrhomboidal than elsewhere, but more those from the several intervening river elongate and inflated, with a tendency for systems. The shells from these rivers tend the ventral margin to be slightly arcuate. to be more solid than those from elsewhere, The periostracum is often rough, black, to be more uniformly biangulate poste- and rayless. The shell form just described riorly, and to have a rather characteristic includes most of the taxa grouped by yellowish brown to shiny chestnut perios- Simpson (1914, 2: 639) under Unio con- tracum, often with fine dark green rays. fertus Lea. In northern Florida, there is an eco- On the Atlantic Slope of Georgia, E. phenotype that lives in lakes and attains icterina (Conrad) is most easily confused much less size than the river one. It has with Elliptio complanata (Lightfoot) with a heavy shell, but tends to be generally which it is associated at many of the same oval. stations, but complanata is quite consist- In the Wekiva River and Oklawaha ently rhomboidal, and the valves are less rivers of the St. Johns River system, inclined to be inflated. The periostracum Florida, there occurs toward the headwaters of icterina is sometimes bright yellow or of the streams and in springs a very thin, chestnut and is generally more shiny and compressed, subrhomboidal ecophenotype smooth than that of complanata. that ends in a broad biangulation below the Range. Apalaehicolan region: Escambia 330 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

River system, east to the St. Marys River Altamaha River Drainage. Georgia: system, Georgia. Peninsular Florida. South- [Altamaha River], Hopeton near Darien, ern Atlantic Slope: Altamaha River system, Mcintosh Co. Frederica, Glynn Co. Georgia, north to the White Oak River, North Carolina. Ogeechee River System

Canoochee River Drainage. Georgia: Specimens Examined Canoochee River, Bulloch Co. Canoochee River, 2 mi. N Claxton, Evans Co. Canoo- St. Marys River System chee River, near mouth, Bryan Co. St. Marys River Drainage. Georgia: (MZUM). St. River, 2 mi. E St. George; St. Marys Ogeechee River Drainage. Georgia: River, Traders Creek, Marys Hill; Spanish Rocky Comfort Creek, 1 mi. N Louisville; W Folkston; St. River, 4 mi. SSE Marys Ogeechee River, 2 mi. S Louisville; Wil- Folkston; all Charleton Co. liamson Swamp Creek, Bartow; Nails Creek, 2 mi. S Bartow; all Jefferson Co. Altamaha River System Barkcamp Creek, 7 mi. E Midville; Buck- Ocmulgee River Drainage. Georgia: head Creek, 14 mi. W Wavnesboro; Buck- Swift Creek, near Macon; Tobesofkee head Creek, 4 mi. E Vidotte; Buckhead Creek, 5 mi. S Macon; both Bibb Co. Creek, 9 mi. SW Waynesboro; Mill Creek, Mossy Creek, 2 mi. S W Kathleen, Houston 5 mi. NE Midville; Rocky Creek, 5 mi. Co. Limestone Creek, Brumbrys Mill, 4 SW Waynesboro; all Burke Co. Ogeechee mi. NE Hawkinsville; Cedar Creek, Foun- River, Scarboro (MZUM); Chew Mill tains Mill, 7 mi. SW Hawkinsville; both Creek, 8 mi. W Millen; Little Buckhead Pulaski Co. House Creek, Bowens Mill, Creek, 1 mi. N Millen; all Jenkins Co. 9 mi. N Fitzgerald; Dicksons Creek, 10 Savannah River mi. NE Fitzgerald; both Ben Hill Co. System Geor- Little Ocmulgee River Drainage. Savannah River Drainage. South Caro- gia: Little Ocmulgee River, 6 mi. NE lina: Abbeville District. (USNM). Turkey 2 Cochran, Beckley Co. Alligator Creek, Creek, 7 mi. NW Edgefield, Edgefield Co. mi. W Alamo, Wheeler Co. Gum Swamp Georgia: Savannah River, 0.5 mi. S Augusta; Creek, 1 mi. N McRae; Little Ocmulgee Little Spirit Creek, DeBruce; both Rich- River, Lumber City (USNM); both Tel- mond Co. South Carolina: Lower Three fair Co. Runs Creek, 8 mi. SW Barnwell, Barnwell Oconee River Drainage. Georgia: Na- Co. Savannah River, 6 mi. W Martin; piers Creek, 6 mi. N Mclntyre, Wilkinson Savannah River, Johnsons Landing, 10 mi. Co. Ford Branch, 4 mi. W Dublin; Turkey W Allendale; both Allendale Co. Georgia: Creek, 9 mi. W Dublin, both Laurens Co. Savannah River, 7.5 mi. NE Shell Bluff; Ohoopee River Drainage. Georgia: Newberry Creek, 2 mi. SE Shell Bluff; Ohoopee River, 4 mi. S Wrightsville; Beaverdam Creek, 7 mi. NW Girard; Ohoopee River, 2 mi. NE Adrian; both Sweetwater Creek, 2 mi. N Girard; all Johnson Co. Ohoopee River, 1 mi. E Burke Co. Brier Creek, 7.5 mi. NE Wrens, Adrian; Mulepen Creek, 2 mi. E Adrian; Jefferson Co. Brier Creek, Chalker Bridge, Ohoopee River, 1 mi. S Norristown; all 6 mi. N Wavnesboro, Burke Co. Brier Emanuel Co. Creek, 6 mi. N Sylvania, Screven Co. Little Ohoopee River Drainage. Geor- Rocky Creek, 6 mi. NW Sardis; Miami gia: Little Ohoopee River, 1 mi. E Kite, Branch, 3 mi. NW Sardis; Mill Race, 2 Johnson Co. Little Ohoopee River, 11 mi. mi. N Sardis; all Burke Co. Near [city of] W Swainsboro, Emanuel Co. Savannah, Chatham Co. (USNM). Atlantic Slope Unionidae • Johnson 331

COMBAHEE RlVER SYSTEM Pedee River Drainage. South Carolina: Lake 7.5 mi. ENE Salkehatchee River Drainage. South Swamp Creek, Oates, Co. Carolina: Salkehatchee River, Broxton Darlington mi. Bridge; Whippy Swamp Creek, 2.5 Waccamaw River System NE Crocketville; both Hampton Co. Waccamaw River Drainage. North Edisto River System Carolina: Lake Waccamaw, [town of] Lake Edisto River Drainage. South Caro- Waccamaw, Columbus Co. South Carolina: lina: Snake Swamp Creek [Orangeburg Ditch, Waccamaw River, 1.25 mi. SE Con- Co. Co.], 6 mi. NE Bamberg, Bamberg Co. N. way, Horry Fork, Edisto River, Orangeburg, Orange- Cape Fear River System burg Co. Edisto River, Canadys Landing, 8 mi. SW St. George; Edisto River, Givhans Deep River Drainage. North Carolina: 2 mi. NW Givhans; both Dorchester Deep River, Gulf (Lea), Chatham Co. Ferry,' Co. Cape Fear River Drainage. North Caro- lina: Rocky River, 11 mi. N Sanford, Chat- Cooper-Santee River System ham Co. Livingston Creek, Columbus Co. Broad River Drainage. North Carolina: Ashe River, Pender Co. Cape Fear River. Green River, 5 mi. ENE Mill Spring, Polk Co. White Oak Ri\t:r System Catawba River Drainage. North Caro- White Oak River Drainage. North lina: Catawba River, Bridgewater, Burke Carolina: White Oak River, Maysville, Co. Catawba River; Long Creek; Beaver Onslow Co. Creek; all Gaston Co. Ashleys Creek; Bissels Pond, Charlotte; both Mecklenburg Elliptio (Elliptio) arctata (Conrad) Co. Plate 10:4-9 Wateree River Drainage. North Caro- Unio arctatus Conrad 1834, Amer. Jour. Sci., 25: lina: Wateree River, 2.5 mi. W Camden; 340, pi. 1, fig. 9 (Black Warrior and Alabama Big Pine Tree Creek, 1.5 mi. S Camden; Rivers [Alabama]; figured type not in ANSP. both Kershaw Co. Lectotype, here selected, ANSP 41356, pi. 10, Alabama from T. A. Santee River Drainage. South Carolina: fig. 4, River, Conrad). Conrad, 1834, New Fresh Water Shells United Poplar Creek, Schulers Fish Pond, near States, p. 36, pi. 5, fig. 4. Santee State Park, Co. Santee Orangeburg Unio strigosus Lea 1840, Proc. Amer. Philos. Soc, Canal; Cooper River (Conrad); near 1: 287 (Chattahoochee River, Columbus [Mus- Charleston; all Berkeley Co. cogee Co.], Georgia). Lea, 1842, Trans. Amer. Philos. Soc, 8: 198, pi. 9, fig. 9; figured holo- Black River System type USNM 85890. Lea, 1842, Obs. Unio, 3: 36. Black River South Carolina: Drainage. Unio tortivus Lea 1840, Proc. Amer. Philos. Soc, Cane Savannah Creek, 4 mi. SW Sumter; 1: 287 (Chattahoochee River, Columbus [Mus- Pocotaligo Creek, 3 mi. S Sumter; [right cogee Co.], Georgia). Lea, 1842, Trans. Amer. Philos. 8: USNM branch] Pocotaligo River, 9 mi. SSE Sum- Soc, 204, pi. 12, fig. 17; 85674 labeled type, does not agree exactly ter; Green Swamp Creek, 2.5 mi. W with the figure and is from Lee Co., Georgia, all Sumter Co. Sumter; Sammy Swamp [Flint River drainage]. Lea, 1842, Obs. Unio, Creek, 3 mi. S Paxville, Clarendon Co. 3: 42. Unio lazarus Lea 1852, Proc. Amer. Philos. Soc, Pedee River System 5: 251 (Abbeville District [Savannah River drainage], South Carolina). Lea, 1852, Trans. Yadkin River Drainage. North Caro- Amer. Philos. Soc, 11: 259, pi. 14, fig. 9; lina: Yadkin River. Stewarts Union Pond, figured holotype USNM 86155. Lea, 1852, Co. Obs. Unio, 5: 15. 332 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Unio perstriatus Lea 1852, Proc. Amer. Philos. short, straight lateral teeth. Right valve Soc, 5: 252 (Abbeville District [Savannah with one low but rather chunky pseudo- River drainage], South Carolina). Lea, 1852, cardinal; one lateral tooth. Beak cavities Trans. Amer. Philos. Soc., 10: 255, pi. 12, shallow, with a few dorsal muscle fig. 3; figured holotype USNM 85892. Lea, very ad- 1852, Obs. Unio, 5: 11. scars under the hinge plate. Anterior Unio gracilentus Lea 1857, Proc. Acad. Nat. Sci. ductor muscle scars well impressed, pos- Gaston Phila., 9: 85 (Catawba River, Co., terior ones less so. Pallial line distinct, North Carolina). Lea, 1862, Tour. Acad. Nat. especially anteriorly. Nacre dirty white or Sci. Phila., ser. 2, 5: 58, pi. 3, fig. 205; figured with slightly holotype USNM 85976. Lea, 1862, Obs. Unio, purplish yellowish splotches, 8: 62. iridescent posteriorly. Unio Lea Proc. Acad. Nat. Sci. perlatus 1863, Length Height Width Phila., 15: 193 (Cape Fear River, Black Rock mm mm mm 2 mi. S Bladen Landing [ = Kings Bluff, Co.], 55 22 14 Alabama River, Alabama. Acad. Nat. North Carolina). Lea, 1866, Jour. Lectotype of Unio arctatus Sci. Phila., ser. 2, 6: 15, pi. 4, fig. 13; figured Conrad. USNM 86006. Lea, 1867, Obs. Unio, holotype 56 24 13 Chattahoochee River, Co- 11: 19. lumbus, Muscogee Co., Clench and EUiptio strigosus (Lea), partim. Georgia. Holotype of Unio Bull. Florida State Mus., 1: 165. Turner, 1956, strigosus Lea. 52 23 12 Abbeville District, South Description. Shell small, seldom exceed- Carolina. Holotype of in ing 60 mm length. Outline elongated, Unio lazarus Lea. arcuate. Valves subelliptical, distinctly 49 25 15 Chattahoochee River, Co- compressed, subsolid, inequilateral. An- lumbus, Muscogee Co., terior end regularly rounded; posterior end Georgia. Figured type of Unio tortivus after more broadly rounded, somewhat biangu- Lea, Lea. late, produced and extending below the base line. Ventral margin arcuate. Dorsal Habitat. Lives in rivers near the shore, margin slightly curved, usually indistinctly among and under rocks. joining the obliquely descending posterior Remarks. On the Atlantic Slope, EUip- margin. Hinge ligament prominent, lo- tio arctata (Conrad) is known only from cated near the middle of the shell. Posterior the several type lots, and from a rather ridge low and rounded, usually with a faint large unlocalized series from the Cape second ridge above. Posterior slope slightly Fear River, North Carolina. In spite of concave and smooth. Umbos not swollen, the very erratic distributional pattern, nor raised above the hinge line, located in there appears to be no morphological dif- the anterior sixth of the shell, their sculp- ference between the specimens from the ture consisting of strong ridges. Disk Cape Fear River and those from the Ala- generally slightly concave, caused by a bama River system, where it is most slight umbonal-ventral sulcus. Surface with abundant. It becomes distinctly rare east- well-marked but delicate growth lines, ward. often smooth and shiny, becoming darker The thinness and compression of the and duller toward the borders. Periostra- valves, subelliptical outline, somewhat pro- cum usually greenish or yellowish with duced posterior end, and quite consistently green rays over the entire surface becom- arcuate ventral margin distinguish this ing wider and more prominent posteriorly, species from any other EUiptio in the sometimes dark burnt-brown especially on Atlantic Slope region. older shells. Range. Alabama-Coosa River system. Left valve with two low, rather vestigial, Apalachicolan region: Escambia River somewhat triangular pseudocardinal teeth. system, east to the Apalachicola River Hinge line short and very narrow; two low, system, but discontinuous. Southern At- Atlantic: Slope Unionidae • Johnson 333

lantic Savannah River Soc, 6: 8, pi. 4, fig. 8 (Head of Chester River Slope: upper system, USNM South Carolina; Catawba River, North [Kent Co.], Maryland; figured holotype 86022). Lea, 1838, Obs. Unio, 2: 8. Carolina of the River Cooper-Santee Unio foliculatus Lea 1838, Trans. Amer. Philos. Fear River system, Lower Cape system. Soc, 6: 38, pi. 11, fig. 33 (Savannah River, North Carolina. [Georgia]; figured holotype USNM 85861). Lea, 1838, Obs. Unio, 2: 38. Specimens Examined Unio duttonianns Lea 1841, Proc Amer. Philos. Soc, 2: 31 (Ogeechee Canal, Savannah [Chat- Savannah River System ham Co.], Georgia). Lea, 1842, Trans. Amer. Philos. Soc, 8: 236, pi. 22, fig. 50; type not in Soutli Caro- Savannah River Drainage. USNM [lost]. Lea, 1842, Obs. Unio, 3: 74. lina: Abbeville District (USNM). Unio sagittiformis Lea 1852, Trans. Amer. Philos. Soc, 10: 277, pi. 22, fig. 35 (Oconee River, Cooper-Santee River System near Athens [Clarke Co.], Georgia; figured holotype USNM 85970). Lea, 1852, Obs. Catawba River Drainage. North Caro- Unio, 5: 33. lina: Catawba River, Gaston Co. (USNM). Unio rostraeformis Lea 1856, Proc. Acad. Nat. Sci. Phila., 8: 262 (Swift Creek, near Macon Cape Fear River System [Bibb Co.], Georgia). Changed to: Unio rostriformis Lea 1858, Jour. Acad. Nat. Sci. Cape Fear River Drainage. North Caro- Phila., ser. 2, 4: 64, pi. 10, fig. 46; probable lina: Fear River, Rlack Rock Land- Cape figured holotype USNM 85973. Lea, 1858, ing, 2 mi. S Kings Bluff, Bladen Co. Obs. Unio, 6: 64. (USNM). Cape Fear River. Unio extensus Lea 1857, Proc. Acad. Nat. Sci. Phila., 9: 31 (Dry Creek, near Columbus [Muscogee Co.], Georgia). Lea, 1858, Jour. Elliptio {Elliptio) lanceolata (Lea) Acad. Nat. Sci. Phila., ser. 2, 4: 68, pi. 12, Plate 10: 10 fig. 49; figured holotype USNM 85995. Lea, Plate 11: 1-6 1858, Obs. Unio, 6: 67. Unio emmonsii Lea 1857, Proc. Acad. Nat. Sci. Obliquaria cuprca Rafinesque 1820, Ann. Gen. Phila., 9: 86 (Roanoke River, Weldon [Hali- Sci. Phys. (Bruxelles), 5: 304, pi. 81, figs. 8, fax Co.], North Carolina). Lea, 1862, Jour. 9 (Le Monongahela et le Potowmak; type not Acad. Nat. Sci. Phila., ser. 2, 5: 56, pi. 2, fig. in ANSP [lost]). Ortmann and Walker (1922: 203; figured holotype USNM 86028. Lea, 31) have pointed out that this species might 1862, Obs. Unio, 8: 60. be either E. dilatata Rafinesque, of the Monon- Unio naviculoid.es Lea 1857, Proc. Acad. Nat. gahela River or producta Conrad [ = lanceo- Sci. Phila., 9: 170 (Buckhead Creek, Burke lata Lea] of the Potomac River. Conrad (1834: both Co.; Macon [Bibb Co.] ; Georgia). Lea, 68), as first reviser of this species, referred it 1858, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: to dilatata. The type locality is here restricted 94, pi. 20, fig. 74; figured holotype USNM to the Monongahela River, Pennsylvania, and 86005, from Buckhead Creek, Lea, 1858, Obs. O. cuprea remains a synonym of E. dilatata Unio, 6: 94. Rafinesque. Unio maconensis Lea 1857, Proc. Acad. Nat. Sci. Unio lanceolatus Lea 1828, Trans. Amer. Philos. Phila., 9: 172 (Flint River, near Macon [Co.], Soc, 3: 266, pi. 3, fig. 2 (Tar River, Tar- Georgia). Lea, 1858, Jour. Acad. Nat. Sci. borough [Edgecombe Co.], North Carolina; Phila., ser. 2, 4: 93, pi. 20, fig. 73; figured figured holotype USNM 85905). Lea, 1834, holotype USNM 86004. Lea, 1858, Obs. Unio, Obs. 1: 8. Unio, 6: 93. Unio Lea Trans. Amer. Philos. angustatus 1831, Unio hazelhurstianus Lea 1858, Proc. Acad. Nat. Soc, 4: 114, pi. 17, fig. 43 (Cooper River, Sci. Phila., 9: 166 (Satilla River, Camden Co., South USNM Carolina; figured holotype 85896; Georgia). Lea, 1859, Tour. Acad. Nat. Sci. South Congaree River, Carolina). Lea, 1834, Phila., ser. 2, 4: 211, pi. 26, fig. 92; figured Obs. Unio, 1: 124. holotype USNM 86009. Lea, 1859, Obs. Unio, Unio productus Conrad 1836, Monography Union- 7: 29. idae, no. 3, p. 31, pi. 14, fig. 1 (Savannah Unio viridulus Lea 1863, Proc Acad. Nat. Sci. River, Augusta [Richmond Co., Georgia]; Phila., 15: 193 (Neuse River, near [6 miles E of] figured holotype ANSP 41397). Raleigh [Wake Co.], North Carolina). Lea, Nat. Sci. ser. 6: Unio fislicrianus Lea 1838, Trans. Amer. Philos. 1866, Jour. Acad. Phila., 2, 334 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

85908. 10, pi. 3, fig. 7; figured holotype USNM posterior point may occasionally be sharp Lea, 1867, Obs. Unio, 11: 14. and turned up above the medial line; Unio nasutilus Lea 1863, Proc. Acad. Nat. Sci. sometimes it below the medial Phila., 15: 192 (Livingston's Creek, Brunswick- drops line, but Co. [NE corner of Columbus Co., Cape Fear more generally it is evenly and broadly River drainage], North Carolina). Changed to: pointed. Ventral margin gently curved, Unio nasutulus Lea Acad. Nat. Sci. 1866, Jour. sometimes almost straight, or a bit arcuate. Phila., ser. 2, 6: 12, pi. 3, fig. 9; figured holo- Dorsal margin straight, forming an indis- tvpe USNM 86027. Lea, 1867, Obs. Unio, tinct 11: 16. angle with the obliquely descending Unio attenuatus Lea 1872, Proc. Acad. Nat. Sci. posterior slope. Hinge ligament rather Phila., 24: 157 (Savannah River; Beaver Creek long and low. Posterior ridge rather [few miles E of Bonaire, Ocmulgee River broadly rounded or subangular, generally drainage], Houston Co.; both Georgia). Lea, with a secondary ridge above it; the 1874, Jour. Acad. Nat. Sci. Phila., ser. 2, 8: 41, ridges sometimes cause the to be pi. 14, fig. 38; figured holotype USNM 85974, point slightly from Beaver Creek. Lea, 1874, Obs. Unio, biangulate. Umbos very low, located in the 13: 45. anterior fourth of the shell, their sculpture Unio rostellum Lea 1872, Proc. Acad. Nat. Sci. consisting of several corrugated, longi- Phila., 24: 160 (Macon [Bibb Co.], Georgia). tudinal ridges. Disk rather flat, or just Lea, 1874, Jour. Acad. Nat. Sci. Phila., ser. concave caused a um- 2, 8: 44, pi. 15, fig. 41; figured holotype slightly by slight USNM 85972. Lea, 1874, Obs. Unio, 13: 48. bonal-ventral sulcus. Surface of the shell Unio exactus Lea 1872, Proc. Acad. Nat. Sci. with numerous concentric striae. Perios- Phila., 24: 159 (Savannah River at Governor tracum fine, subshiny, greenish yellow, Hamilton's [Mansion, 8 mi. S of Hardeeville, olive, often with distinct greenish rays, Jasper Co., South Carolina], Georgia). Lea, when but often 1874, Jour. Acad. Nat. Sci. Phila., ser. 2, 8: 45, esrjecially young, becoming pi. 15, fig. 43; figured holotype USNM 85872. rough and brownish or dirty black with Obs. Lea, 1874, Unio, 13: 49. age. Unio subcylindraceus Lea 1873, Proc. Acad. Nat. Left valve with two stumpy pseudo- Sci. Phila., 25: 422 (Rocky Creek, near Macon cardinal teeth, one in front of the [Bibb Co.]; Carter's Creek [Baldwin Co.]; both other, Georgia). Lea, 1874, lour. Acad. Nat. Sci. both somewhat triangular; the hinder one Phila., ser. 2, 8: 58, pi. 20, fig. 57; figured may be vestigial. Hinge line rather short holotype USNM 85863, from Rocky Creek. and narrow; two long, straight lateral Lea, 1874, Obs. Unio, 13: 62. teeth. Right valve with two Unio arctior fisheropsis De Gregorio 1914, II roughly parallel Naturalista Siciliano, 22: 45, pi. 5, figs. 3a-c pseudocardinals, the posterior one inclined (Ogeecho [Ogeechee] Canal [Chatham Co.], to be triangular, serrated and chunky, the Georgia; type Palermo Mus., Sicily [not seen]). more anterior one, low and vestigial; one Elliptio cupreus (Rafinesque). Ortmann, 1919, lateral tooth. Beak cavities very shallow, Mem. Carnegie Mus., 8: 110, pi. 8, fig. 6. with a few dorsal muscle scars. Elliptio fisherianus (Lea). Ortmann, 1919, Mem. Anterior Carnegie Mus., 8: 113, pi. 8, fig. 7. and posterior adductor muscle scars and pallial line all distinct. Nacre white, Description. Shell generally medium bluish throughout most of its range, usually not white, pinkish, purplish, posteriorly iridescent. exceeding 100 mm in length, though reach- ing over 130 mm in a few localities. Out- Width Lengthmm Heightmm mm line elongated trapezoid, or elongated 34 ovate, lance-head shaped, a little over twice as long as high. Valves usually subcom- pressed, thin to subsolid, occasionally quite solid; inequilateral. Anterior end regularly rounded; posterior end elongated and either sharply or bluntly pointed. The Atlantic Sloik Umonidak • Johnson 335

Anatomy. Discussed and figured by the northern range of the species and again Reardon (1929: 11, pi 5, figs. 1-10). The in the ponds near Wilmington, New Han- glochidia are hookless. over Co., North Carolina. Breeding season. Ortmann (1919: 111) Reardon (1929: 9) notes that there is found gravid females on May 6, June 3-8, no constant distinction either in the anat- 1912, in the Potomac and James Rivers, omy or in the shell morphology. Except Virginia. for their large size, specimens from Ochee Habitat. Lives on sandy bottoms, often Creek, about 5 mi. SE Toomsboro, Wilkin- found crawling about with much of the son Co., Georgia, show the same range of shell out of the sand, also found among variation as do specimens from the rocks and in mud, where the current is not Potomac River, Virginia. too swift. There is much local variation in this Remarks. Elliptio lanceolata (Lea), species throughout its range, and it is easy which ranges from the Escambia River to see how Issac Lea fell to naming all of system of Florida to the Juniata River of the shell forms. The specimens from the the Susquehanna River system of Penn- type locality and from several other of the sylvania, is allied to Elliptio icterina (Con- rivers in North Carolina are usually small, rad) and complanata (Lightfoot) of the shiny, and yellowish, without a trace of Apalachicolan and Atlantic Slope regions. rays, whereas over the rest of its range, While complanata has a subtrapezoidal lanceolata generally has greenish rays. The shape, lanceolata is generally easily dis- species varies considerably as to the ratio tinguished from it by its more elongate, of height to length, the sharpness of the lanceolate shape. Even though the pos- posterior point, whether the point is above terior point may be somewhat rounded, it or below the medial line, and even to the can usually be separated from complanata degree of inflation. It can generally be or icterina because it is over twice as long easily separated from any other species of as high and because the dorsal and ventral Elliptio because it is more than twice as margins are roughly parallel. long as high, with the dorsal and ventral On the Atlantic Slope, E. lanceolata is margins roughly parallel. most easily confused with Ligumia nasuta Also found on the Southern Atlantic (Say), which ranges from the James River. Slope is Elliptio shepardiana (Lea), which Virginia, to the St. Lawrence River, is restricted to the Altamaha River system, Canada. Anatomically they are quite dif- Georgia, and although it is obviously ferent, and the shells of nasuta show sexual closely related to lanceolata, it differs from times as dimorphism and a pattern of green rays lanceolata by being over three that is not found in lanceolata. The male long as high. shells can be confused, but nasuta has a In the Suwannee River system of the silvery white or cream colored nacre, Apalachicolan region and in Peninsular whereas lanceolata may be coppery or Florida, E. lanceolata is replaced by a purple, with more stumpy hinge teeth. The closely related species, Elliptio jayensis females of nasuta have a peculiar expansion (Lea). While jayensis exhibits a range of of the lower margin, rendering them easy variation similar to that of lanceolata, it is to distinguish. generally smaller, with a tendency to be Ortmann (1919: 113) separated fisheri- higher behind. The dorsal and ventral anus from lanceolata with some reluctance, margins are not parallel, and it has more on the basis that fisherianus was slightly numerous darker green rays. more elongated, with a greater taper to the In the Apalachicolan region, it is often posterior part of the shell. A few popu- difficult to separate lanceolata from Ellip- lations of this ecophenotype are found in tio icterina (Conrad), which also some- 336 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

times tends to have a lanceolate shell, but Intyre; Ochee Creek, about 5 mi. SE even so, these strikingly long shells can Toomsboro; Turkey Creek, 4 mi. NE generally be separated from any other Allentown; all Wilkinson Co. Ford Branch, 9 Elliptic) by their height to length ratio. 4 mi. W Dublin; Turkey Creek, mi. W all Elliptic) lanceolata (Lea) seems to be Dublin; Rocky Creek, 8 mi. W Dudley; most abundant on the Atlantic Slope of Laurens Co. Ochwalkee Creek, 2 mi. E Georgia. Here and in the Apalachicolan Glenwood, Wheeler Co. region, individuals reach their greatest size. Ohoopce River Drainage. Georgia: Specimens from the Carolinas and the Ohoopee River, 4 mi. S Wrightsville, John- north are generally smaller and more deli- son Co. Mulepen Creek, 2 mi. E Adrian; cate. Ohoopee River, Norristown; both Emanuel Range. Apalachicolan region: Escambia Co. River system, east to the Apalachicola Little Ohoopee River Drainage. Geor- River system, but discontinuous; Satilla gia: Battleground Creek, 1 mi. N Kite, River system, Georgia. Atlantic Slope: Johnson Co. Altamaha River system, Georgia, north to Ogeechee River System the Juniata River of the Susquehanna River system, Pennsylvania. Ogeeehee River Drainage. Georgia: [Williamson Swamp Creek] Rartow; Nails Specimens Examined Creek, 2 mi. S Bartow; both Jefferson Co. Satilla River System Bark Camp Creek, 7 mi. E Midville; Mill 5 mi. E Midville; Buckhead Creek, Satilla River Drainage. Georgia: Sa- Creek, 9 mi. S all Burke Co. tilla River, Camden Co. Waynesboro; Ogee- chee River, bridge 1.5 mi. SW Oliver, Altamaha River System Screven Co. (MZUM). Ogeechee River, Ocmulgee River Drainage. Georgia: bridge, 1 mi. E Blitchton; Ogeechee River, Walnut Creek, 2 mi. E Macon; Rockv Jinks Bridge [Rte. 16]; Dolly Lake, about Creek (USNM); Swift Creek, 4 mi. E 1.5 mi. below Kiterlighter Camp; all Bryan Macon; Tobesofkee Creek, 5 mi. S Macon; Co. (all MZUM). all Ribb Co. Reaver Creek, E Ronaire Savannah River System (USNM); Mossy Creek, 2 mi. SW Kath- leen; Flat Creek, 2 mi. S Perry; all Houston Savannah River Drainage. Georgia: Co. Ruck Creek, 5 mi. NW Hawkins ville; Savannah River, Augusta, Richmond Co. Limestone Creek, 4.3 mi. E Hawkinsville; (ANSP). Brier Creek, 7.5 mi. NE Wrens, Rig Tuscawhatchee Creek, 5 mi. SW Jefferson Co. Savannah River, 7.5 mi. NE Hawkinsville; Cedar Creek, Fountains Mill, Shell Bluff; Brier Creek, Chalker Bridge, 7 mi. SW Hawkinsville; Mosquito Creek, 6 mi. N Waynesboro; Mill Race, 2 mi. N 8 mi. SE Hawkinsville; all Pulaski Co. Sardis; all Burke Co. South Carolina: House Creek, Rowens Mill, 9 mi. N Fitz- Savannah River, 2 mi. SW Millettville gerald; Dicksons Creek, 10 mi. NE Fitz- ( MZUM ) ; Savannah River, Johnsons Land- gerald; both Ren Hill Co. ing, 10 mi. W Allendale; both Allendale Little Ocmulgee River Drainage. Geor- Co. Georgia: Savannah River, 8 mi. S gia: Little Ocmulgee River, 7 mi. NE Hardeville, Jasper Co. (USNM). Rice Cochran, Reckley Co. Gum Swamp Creek, fields, near Savannah; Ogeechee Canal, 1 mi. N McRae, Telfair Co. Savannah (Lea); both Chatham Co. Oconee River Drainage. Georgia: Oco- Edisto River System nee River, near Athens, Clarke Co. (USNM). Carters Creek, Raldwin Co. Edisto River Drainage. South Carolina: (USNM). Napiers Creek, 6 mi. N Mc- Edisto River [not localized]. Atlantic Slope Unionidae • Johnson 337

Cooper-Santee River System Chowan River System

Cooper River Drainage. Sontli Caro- Nottoway River Drainage. Virginia: lina: Cooper River [not localized]. Nottoway River, 3 mi. E Rawlings, Bruns- Saluda River Drainage. South Caro- wick Co. lina: Saluda River, 2.3 mi. above Ware Shoals, Abbeville Co. James River System Congaree River Drainage. South Caro- James River Drainage. Virginia: Calf- lina: Congaree River, Richland Co. pasture River (Conrad, 1846: 407); North Wateree River South Caro- = Drainage. [ Maury] River, Lexington; [James lina: Wateree River, 2.5 mi. W Camden, River], near Natural Bridge; all Rockbridge Kershaw Co. Co. James River, Buchanan, Botetourt Co. James River, Lynchburg, Campbell Co. Black River System Rivanna River, 2 mi. W Columbia, Flu- vanna Co. Blaek River Drainage. South Carolina: James River, opposite Maidens, Goochland Co. Green Swamp Creek, 2.5 mi. W Sumter; Poeotaligo River, 3 mi. S Sumter; both York River System Sumter Co. South Anna River Drainage. Virginia: Waccamaw River System South Anna River, 1 mi. N. Dabneys, Louisa Co. South Anna River, Gum Tree, Waccamaw River Drainage. South Hanover Co. Carolina: Lake Waccamaw, [town of] Lake Columbus Co. Waccamaw, Rappahannock River System

Cape Fear River System Rapidan River Drainage. Virginia: Rapidan River, N Unionville, Orange Co. Cape Fear River Drainage. North Caro- Branch, N Lignum, Culpeper Co. lina: Livingstons Creek, Brunswick Co. Rappahannoek River Drainage. Vir- (USNM). Wilmington, New Hanover Co. ginia: Rappahannock River, Remington, Co. Water- Neuse River System Faquier Rappahannock River, loo, Culpeper Co. Neuse River Drainage. North Carolina: Little River, Tarpleys Mill, 2 mi. NE Occoquan Creek System Wendell (5); Little River, 2 mi. WSW Oeeoquan Creek Drainage. Virginia: Zebulon (6); both Wake Co. Neuse River, Broad Run [Fairfax Co.] 3 mi. W Manas- 6 mi. E Raleigh (USNM). sas, Prince William Co.

Pamlico River System Potomac River System Tar River Drainage. North Carolina: Potomac River Drainage. West Vir- Tar Tarboro Tar River, (USNM); River, ginia: Back Creek, 0.5 mi. E Glengary, Old 3.5 mi. both Sparta, W Pinetops; Berkeley Co. Shenandoah River, Jefferson Co. Edgecombe Co. Virginia: N Fork Shenandoah River, E Woodstock, Shenandoah Co. Pennsyl- Roanoke Rpver System vania: Rock Creek, 0.75 mi. N Gettysburg, Dan River Drainage. Virginia: Dan Adams Co. Maryland: Aqueduct Lake River [Halifax Co.]. Washington Co. Flat Run, Emmitsburg; both Roanoke River Drainage. Virginia: Monocacv River, 2 mi. W Pleasant; Roanoke River, Weldon, Halifax Co. Frederick Co. Potomac River, Great Falls; 338 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Potomac River, below Alexandria; both more anterior one low and vestigial; one Fairfax Co. lateral tooth. Beak cavities very shallow, with a few dorsal muscle scars. Anterior

Elliptio (Elliptio) shepardiana (Lea) and posterior adductor muscle scars and Plate 11:7 pallial line all distinct. Nacre sometimes bluish white or but Unio shepardianus Lea 1834, Trans. Amer. Philos. pinkish usually pur- or iridescent Soc, 5: 95, pi. 13, fig. 38 [Altamaha River] plish coppery, posteriorly. near Darien [Mcintosh Co.], Hopeton, Georgia; Length Height Width type, not in USNM [lost]; three presumed para- mm mm mm types MCZ 155568 ex Prof. Shepard. Lea, 185 42 26 Altamaha River, 3 mi. NW 1834, Obs. Unio, 1: 207. Simpson, 1914, Cat. Everett City, Glynn Co., Naiades, 2: 698. Georgia. 127 29 15 Ocmulgee River, 1 mi. S Description. Shell often large, reaching Lumber City, Telfair Co., over 185 in Outline mm length. extremely Georgia. elongate, trapezoidal, lance-head shaped, Anatomy. Discussed by Lea (1863: 415). over three times as long as high. Valves Habitat. Lives in banks, or subeompressed to subinflated, subsolid, muddy buried in sand in flowing water. quite inequilateral. Anterior end regularly Remarks. Elliptio shepardiana (Lea) is rounded, usually higher than any other a singular species and it can be confused part of the shell; posterior end very with no other one. It is obviously related elongated and pointed. The point may to E. lanceolata but differs from be at the medial line or below it. Ventral (Lea), it over three times as as margin almost always arcuate. Dorsal mar- by being long high, whereas lanceolata is just over twice gin roughly parallel to the ventral margin, as long as high. In shepardiana the an- long and straight, forming an indistinct terior end is the widest part of the shell; angle with the obliquely descending pos- it is almost alwavs arcuate, and is con- terior margin. Hinge ligament long and sistently pointed. It replaces E. lanceolata low. Posterior ridge rather broadly in the main tributaries of the Altamaha rounded or subangular, generally with a River. secondary ridge above it; the ridges some- Range. Southern Atlantic Slope: re- times cause the point to be slightly bi- stricted to the Altamaha River angulate. Umbos very low, located in the system, Georgia. anterior fifth of the shell, their sculpture of several consisting corrugated, longi- Specimens Examined tudinal ridges. Disk surface flat or just slightly concave caused by a slight um- Altamaha River System bonal-ventral sulcus. Surface of the shell Ocmulgee River Drainage. Georgia: with numerous concentric striae. Perios- Ocmulgee River above Jackson, Butts Co. tracum fine, greenish or with yellowish, Big Tucsawhatchee Creek, 6 mi. S Hawk- faint rays when young, becoming rough insville, Pulaski Co. Ocmulgee River, and brownish or blackish with age. Jacksonville; Ocmulgee River, 1 mi. S Left valve with two stumpy pseudo- Lumber City; both Telfair Co. Oconee cardinal one in front of the teeth, other, River Milledgeville, Baldwin Co. Oconee both somewhat triangular, the hinder one River, 2.5 mi. N Glenwood; Ochwalkee less high. Hinge line rather short and Creek, 2 mi. E Glenwood; both Wheeler narrow; two long straight lateral teeth. Co. Right valve with two roughly parallel Altamaha River Drainage. Georgia: pseudocardinals, the posterior one inclined Altamaha River, 7 mi. N Hazlehurst, Mont- to be triangular, serrated, and chunky, the gomery Co. Altamaha River, 10 mi. NE Atlantic Slope Umonidae • Johnson 339

Surrency, Appling Co. Altamaha River, 22: 169 [probably appeared in April]). There is no 11 mi. N Odum; Altamaha River, "River- way to be sure which name has priority, l)i it Say's name is certainly better known, and side Park," 4 mi. N Jesup; both Wayne Co. it is given primary status here without equi- Altamaha River, 3 mi. NE Everett City, vocation. Co. Altamaha near Fort Rar- Glynn River, Unio dcclivis Say 1831 [1832], Transylvania Jour. rington; Penhollaway Creek, 5 mi. S Fort Med., 4: 527 (Bayou Teche, Louisiana). Say, American Rarrington; Altamaha River, Hopeton, near 1832, Conchology, no. 4 [no pagi- nation], pi. 35; three ANSP 41698 Darien; all Mcintosh Co. syntypes from Mr. Barabino, all smaller than figured type. Genus Uniomerus Conrad Unio camptodon Say 1832, American Conchology, no. 5 [no 42 Uniomcrus Conrad 1853, Proc. Acad. Nat. Sci. pagination], pi. (opposite to New Orleans [Jefferson in Phila., 6: 268. Species listed: U. dcclivis Say, Parish, Louisiana], ponds; type not in ANSP camptodon Say, suhcroccus Conrad, sayii Ward, [lost]). Unio Lea rivularis Conrad, porrectus Conrad, sym- geometricus 1832, Trans. Amer. Philos. 5: metricus Lea, excultus Conrad. Soc, 38, pi. 4, fig. 10 (Bayou Teche, Conrad, 1854, Jour. Acad. Nat. Sci. Phila., ser. Louisiana; figured holotype USNM 85712) 1834, Obs. 1: 150. 2, 2: 296. Lea, Unio, Unio blandin Type species, Unio tetralasmus Say. Subsequent gianns Lea 1834, Trans. Amer. Philos. designation, Simpson, 1900, Proc. United States Soc, 5: 101, pi. 15, fig. 44 (St. Johns River, Natl. Mus., 22: 739. Since U. excultus Conrad Florida; figured holotype USNM 85715). Lea, Obs. 1: 213. is included in Simpson's synonymy of U. tetra- 1834, Unio, lasmus, the subsequent selection of the former Unio declivis Conrad 1836 non Say. See under: by Clench and Turner, 1956, Bull. Florida State Unio rivularis Conrad, 1853. Unio excultus 1 : is Unioni- Mus., 176, invalid under Article 69 ( a ) Conrad, 1838, Monography of Int. dae, no. 11, 1 Orleans ( iv) the Code Zool. Nomen. ( 1964 ). Ort- p. 99, pi. 55, fig. (New mann, 1912a, Ann. Carnegie Mus., 8: 272. [Orleans Parish], Louisiana; type ANSP 20427 [lost]). lists a Frierson (1927: 34-35) number Unio sayii Ward 1839, [in Tappan], Amer. Jour. of species and subspecies under Uniomerus. Sci., 35: 268, pi. 3, fig. 1 (Walnut Creek" and Ohio Canal, near Circleville Like Elliptio, Uniomerus has a wide range [Pickaway Co.], Ohio; type [location unknown]). of environmental tolerance, and while there Unio paralellus [sic] Conrad 1841, Proc. Acad. Nat. are a number of the ecophenotypes, genus Sci. Phila., 1: 20 non Sowerby 1840. Changed appears to be monotypic. to: Unio porrectus Conrad 1845, Jour. Acad. Nat. Sci. ser. 2: 7 Uniomerus fefralasmus (Say) Phila., 2, 296, pi. 26, fig. ([Pearl River] Jackson [Hinds Co.], Mississippi; figured Plate 12: 1-6 holotype ANSP 42847). Unio tetralasmus Say, [September] 1831, American Unio buddianus Lea 1843, Desc. Twelve Uniones Conchology, no. 3 [no pagination] pi. 23 (Lake George and Lake Monroe, Florida). (Bayou St. John [not located] near New Or- Lea, 1845, Trans. Amer. Philos. Soc, 9: 277, leans, Louisiana; type not in ANSP [lost]). pi. 40, fig. 5; figured holotype USNM 85606, Unio obesus Lea 1831, Trans. Amer. Philos. Soc, from Lake George. Lea, 1848, Obs. Unio, 4: 96, 108, pi. 13, fig. 26 (York River, Virginia 4: 35. [corrected to Georgia; Maj. Leconte on p. 108]; Unio symmetricus Lea 1845, Trans. Amer. Philos. figured holotype USNM 85366, labeled, "Little Soc, 10: 73, pi. 4, fig. 11 (Red River, Alex- Ogeechee River [Hancock Co.], Georgia; Maj. andria [Rapides Parish] Louisiana; figured Leconte"). Lea, 1834, Obs. Unio, 1: 106, 118. holotype USNM 85604). Lea, 1848, Obs. Unio, Clench and Turner, 1956, Bull. Florida State 4: 47. Mus., 1: 178, did not see the type, which has Unio paludicolus Gould 1845, Proc. Boston Soc. the corrected locality written on it, and un- Nat. Hist., 2: 53 (Florida Everglades; lecto- fortunately restricted the type locality to Co- type MCZ 169278, selected by Johnson, 1964, lumbus, Georgia. Bull. United States Natl. Mus., 239: 121, pi. Lea, 1854 (Proc. Acad. Nat. Sci. Phila., 7: 243), 31, fig. 3). claimed that this description appeared during Unio ineptus Lea 1852, Trans. Amer. Philos. Soc, the latter part of 1831. It was reported on in 10: 261, pi. 15, fig. 12 (Abbeville District 1832, (Jan. -March number of: Amer. Jour. Sci., [Savannah River drainage], South Carolina; 340 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

figured holotype USNM 85326). Lea, 1852, Unio electrinus Reeve 1865, Conch. Iconica, 16, Obs. Unio, 5: 17. Unio, pi. 25, fig. 121 (Hab. ? Cuming colln; Unio hebes Lea 1852, Trans. Amer. Philos. Soc, type, British Museum (Nat. Hist.) [lost]). Unio Proc. Nat. Sci. 10: 267, pi. 18, fig. 21 (Oconee River, near bisselianus Lea 1867, Acad. Athens [Clarke Co.], Ceorgia; figured holotype Phila., 15: 81 (Bissels Pond, Charlotte [Meck- USNM 85383). Lea, 1852, Obs. Unio, 5: 23. lenburg Co.], North Carolina). Lea, 1868, Unio rivularis Conrad 1853, Proc. Acad. Nat. Sci. Jour. Acad. Nat. Sci. Phila., ser. 2, 6: 277, 85373. Phila., 6: 257. New name for Unio declicis pi. 37, fig. 90; figured holotype USNM Conrad 1836 non Say 1831, in Monography Lea, 1869, Obs. Unio, 12: 37. Sci. Unionidae, no. 5, p. 45, pi. 23, fig. 1 (small Unio jewettii Lea 1867, Proc. Acad. Nat. creek in Green Co., Alabama; figured holotype Phila., 11: 81 (sink of Noonan's [Newnans] ANSP 42852). Lake [Alachua Co.], Florida). Lea, 1868, Jour. Unio paludicolor Conrad 1853, Proc. Acad. Nat. Acad. Nat. Sci. Phila., ser. 2, 6: 276, pi. 37, Sci. Phila., 6: 254. Error for U. paludicolus fig. 89; figured holotype USNM 85374. Lea, Gould. 1869, Obs. Unio, 12: 36. Unio subcroceus Conrad 1854, Jour. Acad. Nat. Unio rivicolus Conrad 1868, Amer. Jour. Conch., 4: 4 near Sci. Phila., ser. 2, 2: 297, pi." 27, fig. 1 (one 280, pi. 18, fig. (brook Tampa of the tributaries to Canadian River, Arkansas; [Hillsborough Co.], Florida; figured holotype type not in ANSP [lost]). ANSP 41411). Unio manubius Could 1855, Proc. Boston Soc. Unio pawensis Lea 1868, Proc. Acad. Nat. Sci. Nat. Hist., 5: 229 (Chihuahua, 60 mi. from Phila., 20: 161 (Paw Creek [Mecklenburg Co.], Gaston Catawba Camp Ringgold = Rio Agualeguas, 3 mi. NE Beaver Co. [ = Creek, Co.], General Trevino, Nuevo Leon [State, Mexico], Run [Gaston and Mecklenburg Cos.]). Lea, teste Taylor, 1967, Veliger, 10: 154; holotype 1868, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: MCZ 169447, figured by Johnson, 1964, Bull. 302, pi. 45, fig. 114; figured holotype USNM United States Natl. Mus., 239: 108, pi. 32, 85380, labeled, "Beaver Creek, [into ?] fig. 5). Catawba Run, North Carolina." Lea, 1869, Unio columbensis Lea 1857, Proc. Acad. Nat. Sci. Obs. Unio, 12: 62. Phila., 9: 31 (Creeks near Columbus [Mus- Uniomerus obesus (Lea). Clench and Turner, cogee Co.], Georgia). Lea, 1858, Jour. Acad. 1956, Bull. Florida State Mus, 1: 177, pi. 5, Nat. Sci. Phila., ser. 2, 4: 75, pi. 14, fig. 55; fig. 2. figured holotype USNM 85360. Lea, 1858, Shell to Obs. Unio, 6: 75. Description, medium large, Unio jamesianus Lea 1857, Proc. Acad. Nat. Sci. reaching 114 mm in length. Outline Phila., 9: 84 ([Pearl River] Jackson [Hinds Co.], rhomboid or long rhomboid. Valves sub- Acad. Nat. Sci. Mississippi). Lea, 1858, Jour. inflated or inflated, subsolid. Anterior end Phila., ser. 2, 4: 53, pi. 6, fig. 35; figured holo- regularly rounded or slightly truncated; type USNM 85365). Lea, 1858, Obs. Unio, 6: 52. posterior end usually somewhat produced.

Unio plantii Lea 1857, Proc. Acad. Nat. Sci. Ventral margin slightly incurved. Dorsal Phila., 9: 171 (Flint River, near Macon [Co.], margin slightly curved, generally forming Lea, 1859, Acad. Nat. Sci. Georgia). Jour. a sharp angle with the almost straight Phila., ser. 2, 4: 192, pi. 21, fig. 76; figured posterior margin. Hinge ligament long holotype USNM 85005. Lea, 1859, Obs. Unio, and located of the 7: 10. [Known only from the holotype, which narrow, posteriorly is a pathological specimen]. umbos. Posterior ridge rounded, ending in Unio cicur Lea 1861, Proc. Acad. Nat. Sci. Phila., a point or feeble biangulation at the base 13: 39 (Little Ocmulgee River, Georgia). Lea, of the shell, sometimes rendering older 1862, Jour. Acad. Nat. Sci. Phila., ser. 2, 5: specimens a bit arcuate. Posterior slope 93, pi. 13, fig. 241; figured holotype USNM often with two radial sulci. low 85532. Lea, 1862, Obs. Unio, 8: 97. Umbos Unio squalidus Lea 1863, Proc. Acad. Nat. Sci. to slightly elevated, located in the an- 15: 192 Neuse near Phila., ( River, Raleigh terior quarter of the shell, their sculpture (Wake Co.]; Roanoke River, near Weldon consisting of five or six heavy ridges that [Halifax Co.]; Deep River; all North Carolina). form a rounded angle on the posterior Lea, 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, ridge, in front of which they tend to be 6: 22, pi. 7, fig. 20; figured holotype USNM black 85376, from Roanoke River. Lea, 1867, Obs. corrugated. Periostracum generally Unio, 11: 26. and slightly roughened, but with a satiny Atlantic Slope Unionxdae • Johnson 341

sheen over most of the surface. Sometimes but like complanata, is generally rayed, the surface is smooth and shiny, especially though often but faintly, whereas tetra- in the umbonal area, and may then be lasmits scarcely ever appears rayed. brownish yellow or yellowish mixed with Uniomerus tetralasmus (Say) is generally green, not rayed. common and abundant throughout the Left valve with two ragged, subequal Apalachicolan region, Peninsular Florida, pseudoeardinal teeth and two straight and in the Southern Atlantic Slope rivers of lateral teeth. Right valve with one tri- Georgia. It becomes noticeably scarce in angular pseudoeardinal often with a vesti- the Carolinas. The periostracum is more gial tooth above it; one lateral tooth. Beak inclined to be smooth on individuals from cavities compressed, but with several this northern end of the range. muscle scars; anterior adductor muscle Range. Interior Basin: Mississippi drain- scars deep, posterior ones faint. Pallial age generally, north to about latitude 40 line distinct. Nacre white, bluish white or degrees, Ohio River. West Gulf Coastal pinkish to lurid purple. region, Alabama-Coosa River system, and Rio Grande River Length Height Width Apalachicolan region: system, Texas, east to the Suwannee River 114 61 39 Ogeeehee River, Bartow, system, Florida. Peninsular Florida. South- Co., Georgia. Jefferson ern Atlantic Slope: Altamaha River system, 102 59 36 Little Cedar 1 mi. E Creek, north to the Nottoway River of the Chowan Wrightsville, Jefferson Co., River svstem, North Carolina. Georgia. 70 40 29 Richardson Creek, 2 mi. Specimens Examined NE Thrift, Jenkins Co., Georgia. Altamaha River System Anatomy. Discussed by Lea (1863: 404). Ocmulgee River Drainage. Georgia: Habitat. lives in smaller Generally Swift Creek, SE Macon, Bibb Co. Mossy streams and on bottoms, ponds muddy Creek, 3.5 mi. SW Kathleen, Houston Co. where it may be locally abundant. Some- = mi. Tucsawhatchee [ Big] Creek, 5 SW times found in sand with com- Elliptic Hawkinsville; Cedar Creek, Fountains Mill, in Southern planata (Lightfoot) larger 7 mi. WSW Hawkinsville; Mosquito Creek, Atlantic Slope rivers, but in this habitat 8 mi. SE Hawkinsville; all Pulaski Co. it is relatively scarce. Brushy Creek, 8 mi. NW Abbeville, Wil- Remarks. In the Apalachicolan and cox Co. House Creek, Bowens Mill, 9 mi. Southern Atlantic Slope regions, Uniomerus N Fitzgerald, Ben Hill Co. tetralasmus (Say) can be confused with Little Ocmulgee River Drainage. Geor- both Elliptio complanata (Lightfoot) and gia: Little Ocmulgee River, 6 mi. NE E. icterina (Conrad). In general, tetralas- Cochran, Bleckley Co. Alligator Creek, 2.5 mits is more inflated than either of them, is mi. SW Alamo, Wheeler Co. Gum Swamp proportionately higher, more acutely angu- Creek, 1 mi. N McRae, Telfair Co. lar where the dorsal margin meets the Oconee River Drainage. Georgia,: Oco- posterior one, and very often has a satiny nee River, near Athens, Clarke Co. periostracum. The yellowish brown, un- (USNM). Black Creek, 7 mi. N Mclntyre; rayed periostracum and the bluish white Napiers Creek, 6 mi. N Mclntyre; Ochee or pinkish nacre of tetralasmus are distinct Creek, about 5 mi. SE Toomsboro; all from the brown, often rayed periostracum Wilkinson Co. Rocky Creek, 6 mi. E and the dark purplish bronze nacre of Dudley; Turkey Creek, 6 mi. W Dublin; complanata. The periostracum of icterina both Laurens Co. Ochwalkee Creek, 2 mi. is sometimes bright yellow or chestnut, E Glenwood, Wheeler Co. 342 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Ohoopee River Drainage. Georgia: ergy Plant, 25 mi. S Augusta (ANSP). Ohoopee River, 4 mi. S Tennible, Wash- South Carolina: Savannah River, 2 mi. ington Co. Little Ohoopee River, 10 mi. SW Millettville (ANSP); Savannah River, NE Wrightsville; Little Cedar Creek, 1 Kingjaw Point, 10 mi. WSW Allendale mi. E Wrightsville; Big Cedar Creek, 1 mi. (ANSP); both Allendale Co. Mouth of SE Wrightsville; outlet of Rollins Pond, 2 mi. Vermezobre Creek, Savannah Wildlife Res- SE Wrightsville; Battleground Creek, 1 mi. ervation, Jasper Co. (MZUM). Georgia: N Kite; Dry Creek, 9 mi. NW Adrian; all Pond near Savannah, Chatham Co. Johnson Co. Pendleton Creek, 8 mi. N Combahee River System Soperton, Treutlen Co. Mulepen Creek, 2 mi. E Adrian; Yam Grande Creek, 3 mi. Salkehatchie River Drainage. South W Swainsboro; Little Ohoopee River, 6 Carolina: Lemon Creek, 5 mi. W Smoaks, mi. SW Swainsboro; all Emanuel Co. Colleton Co. Altamaha River Drainage. Georgia: Edisto River System Hopeton, near Darien, Mcintosh Co. Brunswick, Glynn Co. Edisto River Drainage. South Caro- lina: 20 mi. W Charleston, Charleston Co. Ogeechee River System Edisto River [not localized]. Canoochee River Drainage. Georgia: Cooper-Santee River System Rocky Creek, 1.5 mi. E Swainsboro; Canoo- chee River, 4 mi. E Swainsboro; Little Cooper River Drainage. South Caro- Canoochee Creek, 6 mi. E Swainsboro; all lina: Cooper River (USNM). Emanuel Co. Little Lotts Creek, 2 mi. Catawba River Drainage. North Caro- SE Statesboro; Lotts Creek, 7 mi. SW lina: Beaver Creek, Gaston Co. Paw Creek; Statesboro; both Bulloch Co. Irwins Creek; Bissels Pond, Charlotte; Ogeeehee River Drainage. Georgia: Elias Pond, 10 mi. from Charlotte; all Little Ogeechee River [Hancock Co.] Mecklenburg Co. (USNM). Ogeechee River, Bartow; Nails Wateree River Drainage. South Caro- Creek, 2 mi. S Bartow; Rockv Creek, 2 mi. lina: Wateree River, 2.5 mi. W Camden; Pine 1.5 mi. S S Wadley; all Jefferson Co. Mill Creek, 5 Big Tree Creek, Camden; mi. E Midville; Barkcamp Creek, 7 mi. E both Kershaw Co. Midville; Barkcamp Creek, 9 mi. SW Santee River Drainage. South Caro- lina: mi. Waynesboro; all Burke Co. Spring, Green Swamp Creek, 2.5 W Ogeechee River, 1 mi. S Millen; Richardson Sumter, Sumter Co. Poplar Creek, Schu- lers Fish near Santee State Creek, 2 mi. NE Thrift; both Jenkins Co. Pond, Park, Ogeechee River, Flat Ford Landing Orangeburg Co. Santee River, Cross, (MZUM); Dolly Lake, about 1.5 mi. below Berkeley Co. both Kiterlighter Camp (MZUM); Bryan Black River System Co. Blaek River Drainage. South Carolina: Savannah River System Cowpen Swamp, 11 mi. NE Sumter, Sum- Savannah River Drainage. Soatl} Caro- ter Co. lina: Abbeville District (USNM), Turkey Pedee River System Creek, 7 mi. NW Edgefield, Edgefield Co. Georgia: Beaverdam Creek, 7 mi. NW Lynches River Drainage. South Caro- Girard; outlet of Waynesboro Lake, lina: Lynches River, 2 mi. NE Bishopville, Waynesboro; Mill Race, 2 mi. N Sardis; Lee Co. all Burke Co. Brier Creek, 6 mi. N Sylvania, Yadkin River Drainage. North Caro- Screven Co. Savannah River, Atomic En- lina: Yadkin River. Atlantic Slope Unionidae • Johnson 343

Waccamaw River System Symphynota Simpson 1900, Proc. United States Natl. Mus., 22: 662. Waccamaw River North Drainage. Type species, Symphynota compressa Lea, origi- Carolina: Lake Waccamaw (USNM); nal designation, non Symphynota Lea 1829, the drainage canal beside Lake Waccamaw, type species of which is Unio alata Say. 1 mi. NNW Dupree Landing; both Colum- Original designation. bus Sontli Carolina: Co. Waccamaw River, There is only one species in Lasmigona Wachesaw 2 mi. Murrells Landing, W s. s., L. costata (Rafinesque), which is Inlet, Georgetown Co. found in the Interior Basin and the upper St. Lawrence River system. In addition Cape Fear River System to the subgenera discussed here are Alas- Deep River Drainage. North Carolina: minota Ortmann (Sidcularia Rafinesque Deep River (USNM). is not identifiable, teste Ortmann and Cape Fear River Drainage. North Caro- Walker, 1922: 36) and Pterosyna Rafines- lina: Rocky River, Chatham Co. 11 mi. N que. They do not occur on the Atlantic Sanford, Lee Co. Stream below Greenfield Slope, but belong to the Interior Basin. Mill Pond, Wilmington, New Hanover Co. (ANSP). Subgenus Platynaias Walker Symphynota Simpson 1900, Proc. United States Neuse River System Natl. Mus., 22: 662. [non] Lea. (See under Neuse River Drainage. North Caro- Lasmigona, above.) Platynaias Walker 1918, Occ. Pap. Mus. Zool., lina: Neuse River, near Wake Co. Raleigh, Univ. Mich., 49: 2. Type species, Symphynota compressa Lea, origi- Pamlico River System nal designation. Tar River North Carolina: Drainage. Platynaias probably contains only two, Sandy Creek, Franklin Co. Fishing Creek, probably allopatric species, L. compressa 4 mi. E Leggett; Tar River, Tarboro; both (Lea) (Ortmann 1919: 116, pi. 9, figs. 1, Co. Edgecombe 2 as L. viridis Rafinesque), which is found in the Interior Hudson Roanoke River System Basin, Bay; Upper Mississippi, Ohio, and St. Lawrence River Roanoke River Drainage. North Caro- systems from Saskatchewan and Nebraska lina: Roanoke River, near Weldon, Hali- to Vermont; extending into the Hudson fax Co. (USNM). River on the Atlantic Slope; and L. sub- viridis (Conrad), which is primarily a spe- Chowan RrvER System cies of the Atlantic Slope, but which ex- Nottoway River Drainage. Virginia: tends into the Lake Ontario drainage. Nottoway River. Most of the other taxa listed under this subgenus by Frierson (1927: 20, 21) are Subfamily Anodontinae (Swainson 1840) included elsewhere in this paper in synon- Ortmann 1910 ymy, save for L. neglecta (Lea) and L. are out of the quadrata ( Lea ) which scope Genus Lasmigona Rafinesque of this paper. Subgenus Lasmigona s.s. Rafinesque

Lasmigona Rafinesque 1831, Continuation of Lasmigona (Platynaias) subviridis (Conrad) Monog. Bivalve Shells of River Ohio (Phila.), Plate 12:7-9 p. 4. Species listed: Alasmidonta marginata Plate 13: 1 Say, A. costata Rafinesque. Type species, Alasmidonta costata Rafinesque. Unio subviridis Conrad 1835, New Fresh Water Subsequent designation, Simpson 1900, Proc. Shells United States; appendix, p. 4, pi. 9, fig. United States Natl. Mus., 22: 664. 1 (Schuylkill River; Juniata River [Rlair Co.]; 344 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

figured holotype ANSP 2105 [lost]; creeks in Lasmigona (Platynaias) sidiviridis (Conrad). Lancaster Co.; [all Pennsylvania]). Ortmann, 1919, Mem. Carnegie Mus., 8: 121, Unio viridis Conrad 1836, Monography Unioni- pi. 9, figs. 3, 4. Ortmann and Walker, 1922, Occ Mus. Zool. Univ. no. dae, no. 4, p. 35, pi. 17, fig. 1, non Rafinesque Pap. Michigan, 112, 1820. p. 35. Clarke and Berg, 1959, Cornell Univ. Sta. Mem. no. 31. Unio tappanianus Lea 1838, Trans. Amer. Philos. Exp. 367, p. 32, fig.

Soc, 6: 62, pi. 17, fig. 55 ( [Frankstown Description. Shell generally small, sel- Branch], Juniata River, near Hollidaysburg dom over 55 in Out- [Blair Co.]; figured holotype USNM 85240; reaching mm length. Schuylkill [River]; and a small stream near line subrhomboid or subovate. Valves Obs. Lancaster; [all Pennsylvania] ). Lea, 1838, subeompressed to slightly inflated, subsolid. Unio, 2: 62. Anterior end regularly rounded; posterior Unio hyalinus Lea 1845, Proc. Amer. Philos. Soc, end more broadly rounded and biangulate 4: 164 ([James River drainage], Richmond near the base. Ventral almost [Henrico Co.], Virginia). Lea, 1848, Trans. margin Amer. Philos. Soc, 10: 69, pi. 2, fig. 4; figured straight or slightly curved. Dorsal margin 86131. Obs. holotype USNM Lea, 1848, Unio, straight, sometimes forming a winglike 4: 43. angle with the obliquely descending pos- Unio decoratus Lea 1852, Trans. Amer. Philos. terior margin. Hinge ligament prominent. Soc, 10: 257, pi. 13, fig. 6 (Abbeville District Posterior but [Savannah River drainage] South Carolina; ridge low, double, ending figured holotype USNM 83972). Lea, 1852, near the base of the shell in a slight bi- Obs. Unio, 4: 43. angulation. Posterior slope sometimes Unio Lea Proc. Amer. Philos. pijgmacus 1852, slightly concave above the upper ridge. Soc, 5: 52 (Abbeville District [Savannah River Umbos elevated above the hinge line, drainage] South Carolina). Lea, 1852, Trans. located anterior to the middle of the shell, Amer. Philos. Soc, 10: 262, pi. 15, fig. 14; figured holotype USNM 85240. Lea, 1852, their sculpture consisting of about four Obs. Unio, 5: 18. subnodulous bars, the first two subcon- Unio charlottensis Lea Proc. Acad. Nat. 1863, centric, the others deeply double-looped, Sci. Phila., 15: 191 ( [Sugar Creek], near Char- anterior loops rounded, posterior ones lotte, Mecklenburg Co., North Carolina). Lea, with a few radial threads behind. 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: angular 8, pi. 2, fig. 5; figured holotype USNM 85402. The disk is smooth or slightly roughened Lea, 1867, Obs. Unio, 11: 12. by growth rests; periostraeum yellowish or Unio Lea Proc. Acad. Nat. Sci. pertenuis 1863, brownish green, occasionally olivaceous, Phila., 15: 193 (Neuse River, near [6 mi. E of] subshining to dull, usually faintly rayed Raleigh [Wake Co.], North Carolina). Lea, over the entire the 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: surface, darker on pos- 8, pi. 2, fig. 4; figured holotype USNM 86139. terior slope. Obs. 11: 12. Lea, 1867, Unio, Hinge teeth well developed but rather Unio insolidus Lea 1872, Proc. Acad. Nat. Sci. delicate. Left valve with two lamellate, Phila., 24: 159 (Abbeville District [Savannah that are directed forward River drainage] South Carolina; Fredericksburg pseudocardinals [Spotsylvania Co.], Virginia; Irwin's Creek, and almost parallel to the hinge with a North Mecklenburg Co., Carolina). Lea, 1874, rudimentary interdental projection which Jour. Acad. Nat. Sci. Phila., ser. 2, 8: 40, pi. fits into an interdental groove in the right 13, fig. 37; figured holotype USNM 83974 valve; two long, straight, thin, lateral teeth. from Irwin's Creek, Lea, 1874, Obs. Unio, 13: 44. Right valve with one long pseudoeardinal, Symphynota viridis (Conrad). Simpson, 1914, directed forward, almost parallel to the Cat. Naiades, 1: 484. hinge line; interdentum wide; one thin Symphynota charlottensis (Lea). Simpson, 1914, lateral tooth. Beak cavities shallow, con- Cat. Naiades, 1: 484. taining dorsal muscle scars. Anterior ad- Symphynota decorata (Lea). Simpson, 1914, Cat. ductor muscle scars distinct, posterior ones Naiades, 1 : 486. less so. Pallia] line distinct Unio pygmaeus (Lea). Simpson, 1914, Cat. anteriorly, Naiades, 2: 649. where the shell may be slightly thickened. Atlantic Slope Unionidae • Johnson 345

Nacre bluish white and iridescent, some- Ortmann (1919: 123) calls attention to times pinkish, especially toward the urn- some specimens of L. subviridis from Co- bonal cavities. nococheague Creek, Greencastle, Franklin Width Co., Pennsylvania, remarkable for their Lengthmm Heightmm large size. The largest one measured 63 60 37 19 Abbeville District [Savan- mm in length, 38 mm in height, and 26 nah River drainage], South mm in width. Lea described a single giant Carolina. Holotype of Unio of this decoratus Lea. specimen species under the name Unio from near 30 18 11 As above. Holotype of charlottensis, Sugar Creek, Unio pygmaeus Lea. Charlotte, Mecklenburg Co., North Caro- 116 56 38 Pfeiffers Pond, Charlotte, lina, part of the Catawba River drainage Mecklenburg Co., North of the upper Cooper-Santee River system, Carolina (ANSP). which measured 112 mm in length. During 117 59 29 compressa Lasmigona the last century, a few additional giant (Lea). Small brook, Mid- specimens, as well as a number of smaller dlebury, Addison Co., Ver- mont. ones, were collected in Pfeiffers Pond, Charlotte, Mecklenburg Co., North Caro- Anatomy. Discussed by Ortmann (1911: lina, by C. M. Wheatley (ANSP and 283). Glochidia are subtriangular, almost MCZ). The original label on the lot in with hooks. measure semicircular, They the Academy of Natural Sciences of Phila- 0.36 mm in length, 0.30 mm in height. The delphia (ca. 1900) states that, "the pond host fish is not known. from which they were taken has long since season. This Breeding hermaphroditic- been filled in." is the breeding season species bradytictic, Ortmann (1913a: 371; 1919: 124) found beginning in August and lasting until June L. subviridis to be especially abundant on 1919: 122). (Ortmann, the west side of the Alleghenian divide in Habitat. found in gravel or Usually the Greenbrier and New rivers of the up- bottoms in medium or small streams, sandy per Kanawha drainage in Virginia and as well as canals and the pond areas of West Virginia, where he suggested this streams; seems to avoid large rivers. species originated. Remarks. With its lamellate pseudo- Range. Interior Basin: New and Green- cardinal teeth and characteristic rudiment- brier rivers of the upper Kanawha River ary interdental projection in the left valve, drainage, Virginia and West Virginia. At- Lasmigona subviridis (Conrad) is not con- lantic Slope: upper Savannah River system, fused with other Atlantic any Slope species, South Carolina; north to the Hudson River in New York, where its distribution except system, westward through the Mohawk overlaps that of Lasmigona compressa River and Erie [now New York State in the lower St. Lawrence and (Lea) Barge] Canal to the Genesee River; all Hudson river systems. L. compressa, New York. which is found abundantly in the Ohio- Mississippi drainage of the Interior Basin, Specimens Examined has a rather heavy, compressed, trapezoidal Savannah River System shell, and the left valve has a well-de- Caro- veloped interdental projection. L. sub- Savannah River Drainage. South viridis has almost invariably a smaller, lina: Abbeville District (USNM). thinner, more delicate shell which is more Cooper-Santee River System inflated (see comparative measurements, above), and the interdental projection is Catawba River Drainage. North Caro- rudimentary. lina: Irwins Creek (MZUM); Sugar Creek, 346 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

near Charlotte (USNM); Pfeiffers Pond, River, Buena Vista, Rockbridge Co. (Ort- Charlotte; all Mecklenburg Co. [Ten Mile mann, 1919: 124). Rivanna River, 2 mi. Creek], Wolfsville [7 mi. W Monroe], W Columbia, Fluvanna Co. James River, Union Co. (MZUM). opposite Maidens, Goochland Co. [James River], Richmond, Henrico Co. (Lea). Pedee River System York River System Yadkin River Drainage. North Caro- lina: Crosiers Branch, Cabarrus Co. North Anna River Drainage. Virginia: (ANSP). Church River, Orange Co. (MZUM).

Cape Fear River System Rappahannock River System River North Caro- Cape Fear Drainage. Rapidan River Drainage. Virginia: lina: Cape Fear River, Kinnon, Cumber- Rapidan River, 3 mi. N Orange, Orange land Co. (MZUM). Co. Rapidan River, Rapidan, Culpeper Co. (Ortmann, 1919: 124). Neuse River System Rappahannock River Drainage. Vir- Neuse River North Carolina: Drainage. ginia: Rappahannock River, Remington, Flat 2.5 mi. S Mt. Person River, Tirzah, Fauquier Co. (Ortmann, 1919: 124). [Rap- Co. Flat River, 2.3 mi. ENE (80). Rouge- pahannock River], Fredericksburg, Spot- mont, Durham Co. Eno River, 1.75 (78). sylvania Co. (Lea). mi. ESE Hillsboro Center, Orange Co. (117). Neuse River, 9.5 mi. ENE Durham, Potomac River System Durham Co. (62). Neuse River, 5 mi. Potomac River West Vir- NNW Bayleaf (56); Neuse River, Falls Drainage. ginia: Creek, 8 mi. E (50); Neuse River, 5.75 mi. E Raleigh Sleepy Berkeley Co. (45); Neuse River, 3 mi. S Milburnie (33); Springs, Morgan (MZUM). Virginia: S. Fork, Shenandoah River, Elkton, Rock- Stirrup Iron Creek, 2 mi. NE Morrisville Co. 1919: West (43); Swift Creek, 3 mi. SSW Garner (20); ingham (Ortmann, 124). Little River, Tarpleys Mill, 2 mi. NE Virginia: S. Fork, Shenandoah River, Harp- Wendell (5); all Wake Co. Neuse River, ers Ferry, Jefferson Co. (Ortmann, 1919: 3.5 mi. NNE Smithfield, Johnston Co. (25). 124). Virginia: Potomac River, 5 mi. S Lucketts, Loudoun Co. Potomac River, Pamlico River System Great Falls County Park, Fairfax Co. Tar River Drainage. Nortli Carolina: (USNM). Tar River, Providence, Granville Co. Genus Alasmidonfa Say . mi. ( MZUM ) Tar River, 2 W Springhope, Prolasmidonta Ortmann Nash Co. Subgenus Prolasmidonta Ortmann 1914, Nautilus, 28: 44. Roanoke River System Type species, Unio heterodon Lea, original designation. This subgenus is monotypic. Roanoke River Drainage. Virginia: Stone River, Clarksville, Mecklenburg Co. Alasmidonfa heterodon (Lea) is restricted (MZUM). to the Atlantic Slope region. The general shape and the distinct posterior ridge give River System James the shell the unmistakable look of an Appomattox River Drainage. Virginia: Alasmidonfa, but it is not close to the other Appomattox River, Petersburg, Dinwiddie members of the genus. There are two Co. (MZUM). lateral teeth in the right valve and one in James River Drainage. Virginia: North the left which is the reverse arrangement Atlantic Slope Unionidae • Johnson 347

of these teeth in all other Atlantic Slope disk is smooth; the periostracum is green- Unionidae. ish olive to brownish. In addition to in- distinct concentric bands of lighter and heterodon Alasmidonta (Prolasmidonta) darker green, there are often obscure green (Lea) rays which are occasionally sharp and well Plate 13:2 defined. teeth delicate. Left valve with Unio heterodon Lea 1830, Trans. Amer. Philos. Hinge Soc, 3: 428, pi. 8, fig. 11 (Schuylkill [River] two lamellate pseudocardinal teeth which and Derby Creek [mostly in Delaware Co.], are parallel to the hinge, the anterior one to Mr. Pennsylvania; figured type, belonged sometimes rudimentary; an interdental Mason, not in USNM or ANSP [presumed lost]. tooth that may be well developed and iso- Lea mentions specimens in possession of T. A. lated or connected with the Conrad, now ANSP 41004, 41005). Lea, 1834, posterior Obs. Unio, 1: 42. pseudocardinal; one lateral tooth. Right Alasmidonta heterodon (Lea) 1914, Simpson, Cat. valve with two compressed crenulated 1: 499. Naiades, pseudocardinals, the more anterior one . \lasmidonta ( Prolasmidonta ) heterodon ( Lea ) . often rudimentary; interdentum narrow Ortmann, 1919, Mem. Carnegie Mus., 8: 173, and short; two delicate lateral the pi. 12, figs. 1, 2. Clarke and Berg, 1959, teeth, Cornell Univ. Exp. Sta. Mem. no. 367, p. 24, upper one often partially rudimentary. fig. 35. Beak cavities shallow, containing dorsal muscle scars. Anterior adductor muscle Description. Shell small, generally from scars ones less so. Pallial 30 to 45 mm, seldom reaching more than distinct, posterior line distinct Nacre bluish 50 mm in length. Outline subrhomboidal anteriorly only. or white, often cream colored or or subtrapezoidal, sometimes more or less silvery yellowish toward the umbonal cavities. elongate. Valves rather flat, though fe- Sexual differences are visible in the shell. males are somewhat swollen in the region In the male shell is compressed, of the posterior ridge, thin but strong. general ovate, and with the ventral mar- Anterior end regularly rounded; posterior elongate, curved, the ridge not sharp, end somewhat produced and angular. gin posterior and the slope not truncate, Ventral margin straight or slightly arcuate. posterior whereas the female shell is swollen in the Dorsal margin short and straight, almost of the always forming a sharp angle with the region posterior ridge, rendering the truncate, the outline of obliquely descending posterior margin. posterior slope the shell more and the ventral Hinge ligament occupying almost all of the trapezoidal, concave. posterior dorsal margin. Posterior ridge margin slightly distinct though rounded, some- Length Height Width broadly mm mm mm times almost angular in females, usually 52 26 17 [Ten Mile River], ML\- ending in a broad point near the base. ville, Cheshire, New Haven Posterior or slope flat, slightly concave, Co., Connecticut. Male. with one or two fine lines. radiating Umbos 40 23 16 Neuse River, Pooles Bridge, somewhat inflated but not very prominent, Wake Co., North Carolina. located slightly anterior to the middle of Male. Little 2 mi. NE the shell, their sculpture consisting of three 38 22 18 River, Wendell, Wake Co., North or four, sometimes five, bars, the first two Carolina. Female. concentric, the others with a distinct angle on the posterior ridge in front of which is Anatomy. Discussed by Lea (1863: 442) a shallow sinus. The bars are rather heavy; and Ortmann (1911: 295). Glochidia are the sinus does not assume the shape of a subtriangular with strong hooks. They re-entering angle, and the sculpture can measure 0.30 mm in length, 0.25 mm in not be regarded as double-looped. The height. The host fish is unknown. 34S Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Breeding season. Ortmann (1919: 174) Rappahannock River System reports gravid females having been taken Rapidan River Drainage. Virginia: in and It is February April. bradytictic. Blue Run, Orange Co. (MZUM). Habitat. Lives in water that does not Rappahannock River Drainage. Vir- flow very fast, in gravel, sand, or in muddy ginia: Mountain Run, Culpeper Co. (Ort- sand. mann, 1919: 175). Remarks. Alasmidonta heterodon (Lea) is an Atlantic Slope species, not closely Potomac River System related to the other members of the genus. Potomac River Drainage. Maryland: It can be confused with no other unionid Mcintosh Run, 4 mi. N Leonardtown, St. in the region because of its distinct den- Marys Co. (H. D. Athearn). tition. A. heterodon has two lateral teeth in the right valve, though the upper one Subgenus Alasmidonta s.s. Say may be rather rudimentary, and one in the Monodonta Say 1817, Nicholson's Encyclopedia, left valve. This tooth arrangement is just 2 [no pagination] non Lamarck 1799. the of all other Atlantic opposite Slope Alasmidonta Say 1818, lour. Acad. Nat. Sci. Unionidae. Phila., ser. 1, 1: 459. This species is usually rare, and its er- Type species, Monodonta undulata Say. Mono- ratic distribution may be merely a reflec- typic. Alasmodonta [sic] Say 1819, Nicholson's Encyclo- tion of inadequate collecting. Judging from pedia, ed. 3, 4 [no pagination]. the available it to be most records, appears Hemiodon Swainson 1849, Treatise on Malacology, widely distributed and abundant in the pp. 288, 381. Species listed: H. undulatus Connecticut River system of Vermont, New [Say], An. rugosus, purpurascens, arcolata non Hemiodon Hampshire, Massachusetts, and Connecti- [all Swainson], Swainson, op. cit., p. 191. cut. Type species, A. undulata Say. Subsequent Range. Atlantic Slope: Neuse River designation, Hermannsen, 1847, Indicis Gene- system, North Carolina, north to the rum Malacozoorum, 1: 527. Swainson Treatise on Petitcodiac River system, New Brunswick, Unio))sis 1840, Malacology, pp. 289, 382. Canada, but discontinuous. Known only Type species, U. mytiloides Swainson and U. from the additional following intervening radiata Swainson. Both names are based on River drainage systems; Rappahannock the same fig., no. 64 = Alasmidonta undulata system, Virginia; Potomac River system, (Say). BuUella Proc. United States Natl. Maryland; Delaware River system, Penn- Simpson 1900, Mus., 22: 672. sylvania; Housatonic River system, Con- Type species, Margaritana arcida Lea. Original necticut (Linsley, 1845); Quinnipiac River designation. system, Connecticut; Connecticut River sys- Jugosus Simpson 1914, Cat. Naiades, 1: 357. tem, Vermont, New Hampshire, Massa- Type species, Strophitus icrightianus Walker. Original designation, Simpson, 1900, Proc. chusetts, Connecticut; Merrimac River sys- United States Natl. Mus., 22: 666. Ortmann, tem, Massachusetts. 1914, Nautilus, 28: 45.

Specimens Examined Alasmidonta s. s. is restricted to the Apalachicolan and Atlantic Slope regions. Neuse River System There is one other species in this subgenus Neuse River Drainage. Nortli Carolina: that is not covered in this paper, A. icright- Neuse River, Pooles Bridge [not located]; iana (Walker) (Johnson, 1967: 8, pi. 2, Neuse River [6 mi. E Raleigh]; Little fig. 5), from the Ochlockonee River system, River, Tarplcys Mill, 2 mi. NE Wendell Florida. (5); all Wake Co. Chicod Creek, Pitt Co. In addition to the subgenera discussed (MZUM). here, or placed in synonymy above, Frier- Atlantic Slopk Umonidae • Johnson 349

son (1927: IS, 19) includes in his list two end regularly rounded; posterior end rather other subgenera of Alasmidonta, Piesso- broadly pointed. Ventral margin almost donta Simpson, which appears to be repre- always slightly rounded. Dorsal margin sented in the Interior Basin by a single short and straight, forming an angle with species, A. calccola (Lea); and Sulcularia the obliquely descending posterior margin. Rafinesque, which Ortmann and Walker Hinge ligament located posteriorly of the (1922: 33) show is not identifiable. The umbos, occupying most of the dorsal mar- species listed under it belong in the genus gin. Posterior ridge rounded, sometimes Lasmigona, subgenus Alasminota Ortmann. with a faint second ridge above it. Posterior slope slightly concave, sometimes with Alasmidonta (Alasmidonta) undulata (Say) irregular oblique corrugations. Umbos Plate 13:3-4 rather inflated and somewhat raised above the hinge line, located anterior to Unto undulata Say 1817, Nicholson's Encyclo- the middle of the shell, their sculpture pedia, 2 [no pagination], pi. 3, tig. 3 (Dela- of four or five ware and Schuylkill rivers [near Philadelphia, consisting strong ridges Philadelphia Co., Pennsylvania]; type, not in running nearly parallel to the growth lines. ANSP [lost]). These ridges extend some distance on the Unto Genera of Recent glabratus Sowerby 1823, disk, and posteriorly are angular, thickened, and Fossil no. 16, 3 (no locality, Shells, fig. and raised. They are crossed by fine, type not in British Museum (Nat. Hist.) [lost]). wrinkled, threads. In front of non Lamarck 1819. radiating and behind the regular umbonal sculpture Alasmidonta sculptilis Say 1829, New Harmony there are lirae. the en- [Indiana] Disseminator, 2, no. 22, p. 339 (Vir- radiating Generally ginia; type not in ANSP [lost]). tire surface of the shell is smooth, though Valenciennes in Humboldt and Unto hians 1827, occasionally slightly roughened, especially Bonpland, Voyage aux Regions Equinoxiales du posteriorly; periostracum greenish, yellow- Nouveau Continent, Pt. 2, 2: 235, pi. 54, figs. ish, reddish brown, with more or less 2a, b (environs de Philadelphie [Philadelphia distinct to blackish of Co., Pennsylvania]; type [location unknown]). green rays varying Uniopsis radiata Swainson 1840, Treatise on width over the entire surface. The pos- Malacology, p. 289, fig. 62 ([North America] terior slope is often somewhat lighter than type [location unknown]). the rest of the shell, with finer and sharper Uniopsis mytiloides Swainson 1840, Treatise on rays. Older shells become blackish, but Malacology, p. 382, fig. 62 ( [North America] the are still visible trans- type [location unknown]. [Both this and the rays through preceding were found on the same figure.] mitted light. Unio swainsoni Sowerby 1868, Conch. Iconica, Left valve with a compressed, sometimes 396 Hab. ? holo- 16, Unto, pi. 76, fig. ( figured stumpy pseudocardinal tooth, often with British Museum [Nat. Hist.] 1900.3.19.21.). type a rudimentary one in front; interdentum Alasmidonta undulata (Say). Simpson, 1914, Cat. rather broad, occasionally with a visible Naiades, 1: 494. but connected interdental tooth; lateral Alasmidonta (Alasmidonta) undulata (Say) Ort- valve with one mann 1919, Mem. Carnegie Mus., 8: 117, pi. tooth barely distinct. Right 11, fig. 7. Clarke and Berg, 1959, Cornell rather high, chunky, subtriangular tooth; no. 33. Univ. Exp. Sta. Mem. 367, p. 25, fig. often with a broad interdentum; with just and Natl. Mus. Athearn Clarke, 1962, Canada, a trace of a lateral tooth. Beak cavities Bull. 183, p. 24, pi. 3, figs. 3, 4. moderately deep with muscle scars under Description. Shell generally small, from the hinge plate. Anterior adductor muscle 50 to 65 mm, though sometimes reaching scars deep, posterior ones faint. Pallial 80 mm in length. Outline elliptical or sub- line distinct anteriorly where the shell is rhomboid. Valves somewhat inflated, thin, thickened. Nacre white, salmon pink, or becoming quite solid anteriorly. Anterior reddish, iridescent posteriorly. 350 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Sexual differences are not visible in the Specimens Examined shell. Cooper-Santee River System Length Width mm Heightmm mm Catawba River Drainage. Nortli Caro- lina: Catawba River 80 51 39 Tar River, Bruce, 9 mi. (USNM). NW Pitt Co., Greenville, Pedee River System North Carolina. 72 45 31 Neuse River, Falls, Wake Yadkin River Drainage. North Caro- Co., North Carolina. lina: Uwhairie River (USNM). 54 32 23 Nottoway River, 3 mi. E Rawlings, Brunswick Co., Cape Fear River System Virginia. Cape Fear River Drainage. North Caro- Anatomy. Discussed by Ortmann (1911: lina: Morgan Creek, 1 mi. SE Chapel Hill, 296). Glochidia are moderately large with Orange Co. (MZUM). Rocky River, 11 strong hooks. They measure 0.34 mm in mi. N Sanford, Chatham Co. (MZUM).

and 0.36 in fish Fear River . length mm height. The host Cape ( MZUM ) is unknown. Neuse River System Breeding season. Ortmann (1919: 178) states that this species is clearly bradytietic, Neuse River Drainage. North Carolina: with a short interim in June and July. Neuse River, Falls (50); Neuse River, Habitat. Lives in water that does not 5.75 mi. E Raleigh Center (45); Swift Creek, flow very fast, in gravel or sand, appears 3 mi. SSW Garner (20); Middle Creek, to avoid mud. 6.25 mi. SE Apex (18); Little River, Tar- Remarks. Alasmidonta undulata (Say), pleys Mill, 2 mi. NE Wendell (5); all of the Atlantic Slope, is not closely related Wake Co. to any Alasmidonta of the Interior Basin. Pamlico River System Its nearest relatives are A. arcula of the Altamaha River system, Georgia, and A. Tar River Drainage. North Carolina: triangulata of the Apalachicolan and South- Tar River, Old Sparta, 3.5 mi. W Pinetops, ern Atlantic Slope regions; but both of Edgecomb Co. Tar River, Bruce, 9 mi. these species are more triangular in outline, NW Greenville, Pitt Co. with shaqoer posterior ridges. A. undulata Chowan River System also resembles A. wiightiana (Walker) of the Ochlockonee River system, Florida, in Nottoway River Drainage. Virginia: shape, and it has a similar tooth structure Nottoway River, 3 mi. E Rawlings, Bruns- and rounded posterior ridge, but wrighb- wick Co. iuna has strong ridges which cover the James River System posterior slope, whereas undulata is un- sculptured. James River Drainage. Virginia: Calf- A. undulata is a common species, espe- pasture River (Conrad, 1846: 407); North on the = cially Northern Atlantic Slope, and [ Maury] River, 2 mi. WNW Lexington is often locally abundant. It seems to be (MZUM); North River, Buena Vista somewhat less common on the Southern (USNM); all Rockbridge Co. Rivanna Atlantic Slope. It avoids the larger rivers, River, 2 mi. W Columbia, Fluvanna Co. preferring smaller streams, in which it James River, opposite Maidens, Goochland often goes far up toward the headwaters. Co. Range. Atlantic Slope: Catawba River, York River System North Carolina, of the upper Cooper- Santee River system, north to the lower St. North Anna River Drainage. Virginia: Lawrence River system, Canada. Mine Run, Orange Co. (MZUM). Atlantic Slope Unionidm: • Johnson 351

Rappahannock River System Cat. Naiades, 1: 509. Clench and Turner, 1956, Bull. Florida State River Mus., 1: 180, pi. 5, fig. 4. Rapidan Drainage. Virginia: Strophitus wrightianus Clench and Turner 1956, Blue Run (MZUM); Mountain Run Bull. Florida State Mus., 1: 180 non Walker (MZUM); both Orange Co. 1901, Nautilus, 15: 65, pi. 3. Rappahannock River Drainage. Vir- Description. Shell generally small, though ginia: Rappahannock River, Remington; reaching 70 mm in length in the Apa- Marsh Run, Remington (both Ortmann, lachieola River system. Outline subtri- 1919: ISO); both Fauquier Co. angular. Valves inflated, thin to rather solid. Anterior end Occoquan Creek System regularly rounded; posterior end rather broadly pointed. Ven- Creek Occoquan Drainage. Virginia: tral margin almost always slightly rounded. Broad Run [Fairfax 3 mi. Ma- Co.], W Dorsal margin short and straight, forming nassas, Prince William Co. an angle with the obliquely descending posterior margin. Hinge ligament occupy- Potomac River System ing almost all of the posterior dorsal mar- Potomac River Drainage. Maryland: gin. Posterior ridge high and angular with Wills Creek, Ellerslie, Allegany Co. (Ort- a second or third faint ridge above it, the mann, 1919: 179). Potomac River, Han- second one ending in a slight biangulation cock, Washington Co. (Ortmann, 1919: near the base of the shell. Umbos full and 179). Pennsylvania: Great Tonoloway high, located anterior to the center of the Creek, Thompson Township, Fulton Co. shell, their sculpture consisting of a number (Ortmann, 1919: 179). Conococheague of strong ridges running nearly parallel to Creek, Greencastle and Scotland, Franklin the growth lines. These ridges extend some Co. (Ortmann, 1919: 179). Virginia: distance on the disk and are crossed by N Fork, Shenandoah River, Broadway, fine, wrinkled, radiating threads. In front Rockingham Co. (Ortmann, 1919: 179). of and behind the regular umbonal sculp- S River of S Fork, Shenandoah River, ture there are radiating lirae. The disk is Waynesboro, Augusta Co. (Ortmann, 1919: smooth; the periostraeum is usually fine 179). S Fork, Shenandoah River, Elkton, and limited to the posterior slope. Young Rockingham Co. Ortmann, 1919: 179). shells may be yellowish or greenish yellow, West Virginia: Shenandoah River, Harpers often with dark green rays of varying Ferry, Jefferson Co. (USNM). Virginia: width. Older shells become dark brownish Potomac River, Great Fairfax Co. Falls, black, but the rays and yellowish ground color are still visible when seen through Alasmidonta (Alasmidonta) triangulata transmitted light. (Lea) Left valve with a compressed, sometimes 91016 13:5,6 stumpy pseudocardinal tooth, often with Margaritana triangulata Lea 1858, Proc. Acad. a rudimentary one in front; interdentum Nat. Sci. 10: 138 Chattahoochee Phila., (Upper broad, occasionally with a visible but con- [River], Georgia; Columbus, Georgia; Polato nected interdental tooth; lateral tooth Co. [sic] [Potato Creek, Upson Co.], Georgia; Sawney's Creek [about 8 mi. NW Camden, barely visible. Right valve with one rather Kershaw Co.], South Carolina). Lea, 1859, high subtriangular tooth; broad interden- Acad. Nat. Sci. ser. 4: Jour. Phila., 2, 228, pi. tum; lateral tooth barely visible. Beak 32, fig. Ill; figured holotype USNM 86249, cavities deep, with dorsal muscle scars from Upper Chattahoochee [River], Lea, 1859, under the Obs. Unio, 7: 46. hinge plate. Anterior adductor Alasmidonta triangulata (Lea). Simpson, 1914, muscle scars deep, posterior ones faint. 352 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Pallial line distinct anteriorly where the Specimens Examined shell is thickened. Nacre bluish white to Ogeechee River System salmon pink. Sexual differences are not visible in the Ogeechee River Drainage. Georgia: shell. Ogeechee River, Scarboro (MZUM); Ogeechee River, bridge, 1 mi. S Dover; Width Length Height both Screven Co. (MZUM). 68 47 38 Savannah River [Johnsons Savannah River System Landing], 10 mi. W Allen- South dale, Allendale Co., Savannah River Drainage. South Caro- Carolina (MZUM). lina: Savannah River [Johnsons Landing], 48 32 27 Mill 2 mi. N Raee, Sardis, 10 mi. W Allendale, Allendale Co. Burke Co., Georgia. (MZUM). Georgia: Mill Race, 2 mi. N Burke Co. Habitat. Prefers sandy mud in rocky Sardis, and eddies of creeks and rivers. pools large Cooper-Santee River System Remarks. Alasmidonta triangulata (Lea) Wateree River South Caro- of the Apalachicolan and Southern Atlantic Drainage. lina: Sawney's Creek [about 8 mi. NW Slope regions cannot be confused with A. River Camden, Kershaw Co.] (USNM). arcula ( Lea ) of the Altamaha system because it is proportionally not as high. Alasmidonta (Alasmidonta) arcula (Lea) The umbos are anterior to the center, and Plate 13: 7, 8 while incurved, do not render the hinge arcula Lea Naiades, line concave, and the posterior ridge is not Margaritana 1836, Synopsis p. 43 [nomen nudum]. Lea, 1838, Trans. Amer. as acutely sharp. Philos. Soc., 6: 71, pi. 22, fig. 69 (Altamaha is closest to A. A. triangulata undulata [River], Liberty [now Long] Co., Georgia; (Say) of the Atlantic Slope region, but the figured holotype USNM 86170). Lea, 1838, Obs. 2: 71. latter has a shape which is usually an Unio, Alasmidonta arcula (Lea). Simpson, 1914, Cat. elongated oval, a very broadly rounded Naiades, 1: 508. posterior ridge, more ponderous pseudo- cardinals, and more of an interdentum. In Description. Shell medium, reaching 75 in Outline Valves the Apalachicolan region, A. triangulata mm length. triangular. much thin but Anterior can be confused only with A. wrightiana inflated, strong. end rounded; end (Walker); the latter has the shape, regularly posterior truncated. Ventral margin straight rounded posterior ridge, and tooth struc- sharply or slightly rounded. Dorsal margin short ture of A. undulata, differing from un- and curved, forming a sharp angle with dulata and triangulata in the strong ridges the obliquely truncated posterior margin. which cover the posterior slope. Hinge ligament occupying almost all of A. triangulata is a rare species through- the posterior dorsal margin. Posterior out its range, but is especially so on the ridge high and sharply angular, with a sec- Southern Atlantic Slope where each of the ond faint ridge above, ending in a slight lots examined consists of one or two only biangulation near the base of the shell. specimens. Posterior slope very truncated. Umbos restricted Range. Apalachicolan region: extremely full and high, located at the to the Apalachicola River system. Southern center of the shell, their sculpture consist- Atlantic Slope: Ogeechec and Savannah ing of a number of strong ridges running river systems, Georgia; and Cooper-Santee nearly parallel to the growth lines. These River system, South Carolina. ridges extend well out on the disk and Atlantic Slope Unionidae • Johnson 353

posteriorly are angular, considerably thick- posterior ridge which forms almost a 90° ened, and raised. They are crossed by angle with the posterior slope. fine, wrinkled, radiating threads. In front This is a rare species; not more than a of and behind the regular umbonal sculp- few specimens have ever been collected at ture there are radiating lirae. The disk is any station in this century save for a series smooth; the periostracum is usually fine of twenty-two specimens collected by H. and limited to the posterior slope. Shell D. Athearn in 1962 in the Ocmulgee River, dull greenish yellow with distinct green below Lumber City, Telfair Co., Georgia. rays of varying length over the entire sur- The old unlocalized specimens of areata face. Mature shells are often blackish, but in the major museums are presumed to the rays are still visible through trans- have been collected near the type locality mitted light. in the tidewater region of the Altamaha Left valve with a long compressed River, where it may be, or was, more pseudocardinal tooth, often with a rudi- abundant. mentary one in front; interdental tooth Range. Southern Atlantic Slope: re- distinct but connected; lateral teeth barely stricted to the Altamaha River system, visible. Right valve with one compressed, Georgia. triangular tooth; scarcely any interdentum; lateral tooth barely visible. Beak cavities Specimens Examined with dorsal muscle scars under very deep, Altamaha River System the hinge plate. Anterior adductor muscle River scars deep, posterior ones faint. Pallial Ocmulgee Drainage. Georgia: Limestone line distinct anteriorly where the shell is Ocmulgee River, Hawkinsville; thickened. Nacre bluish white or white. Creek, near Hartford (H. D. Athearn); Sexual differences are not visible in the both Pulaski Co. Ocmulgee River, below Telfair Co. shell. Lumber City, (H. D. Athearn). Ohoopee River Drainage. Georgia: Length Height Width mm mm mm Ohoopee River, 3 mi. N Leman, Emanuel Co. 80 60 50 Altamaha River [Long Co.], Georgia. Altamaha River Drainage. Georgia: 67 51 44 Ocmulgee River, Hawkins- Altamaha [River], Liberty [now Long] Co. Pulaski ville, Co., Georgia. (USNM). Altamaha River, Fort Barring- 50 43 38 Altamaha River, 4 mi. NE ton, Mcintosh Co. (ANSP). Jesup, Wayne Co., Geor- gia. Subgenus Decurambis Rafinesque Anatomy. Discussed by Lea (1863: 447). Decurambis Rafinesque 1831, Continuation of Habitat. Lives in mud below sand sandy Monograph Rivalve Shells of River Ohio bars in water and eddies. sluggish (Phila. ), pp. 4, 5. Species listed: Alasmodon Remarks. Alasmidonta arcula (Lea) of scriptum Rafinesque and A. atropurpureum the Altamaha River system cannot be con- Rafinesque. Alasmodon = fused with any other unionid in the At- Type species, scriptum Rafinesque Alasmidonta marginata (Say). Subsequent lantic Slope region, it is close to though designation, Ortmann and Walker, 1922, Occ. A. which is found in triangulata (Lea), Pap. Mus. Zool., Univ. Mich., no. 112, p. 38. both the Apalachicolan and Atlantic Slope Rugifera Simpson 1900, Proc. United States Natl. regions. Alasmidonta arcula differs from Mus., 22: 670. Alasmidonta triangulata by being proportionally higher Type species, marginata Say. Original designation. with centrally located umbos which are so elevated as to be incurved, by its concave The present author follows Ortmann and hinge line, and by its extremely sharp Walker (1922: 38) and regards Decuram- 354 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

bis as a subgenus of Alasmidonta, though rounded, with a system of fine radial Fricrson (1927: 21) raised Decurambis to wrinkles running to the upper posterior a genus. margin. Umbos large and somewhat in- Decurambis includes but two allopatric flated, but not high, located in the anterior species: A. marginata Say (Ortmann, 1919: third of the shell, their sculpture consisting three thick bars. disk is LSI, pi. 12, fig. 3), primarily a species of of or four The the Interior Basin, but which is also found smooth; the periostraeum is yellowish, on the Northern Atlantic Slope in the Sus- greenish, brownish, or blackish, generally quehanna and upper St. Lawrence drainage with distinct narrow or wide, straight, systems; and A. varicosa (Lamarck), which greenish or blackish rays. The rays are is limited in distribution to the Atlantic- usually broken by growth lines. Slope. Each valve has one thin, distinct, de- pressed, triangular pseudocardinal; no Alasmidonfa {Decurambis) varicosa lateral teeth. Beak cavities moderately (Lamarck) deep with muscle scars under the hinge Plate 14: 1,2 plate; anterior adductor muscle scars deep, Unio varicosa Lamarck 1819, Hist. Nat. des posterior ones less so. Pallia] line distinct Animaux sans 6: 78 riviere de Vertebres, (la anteriorly where the shell is slightly thick- de Philadelphie Schuglkill [Schuylkill] pies ened. Nacre bluish white, often with [Philadelphia Co., Pennsylvania]; holotype, or shades. Geneva Museum, teste Johnson, 1953, Nautilus, salmon, pinkish, purplish 66: 95; aussi dans le lac Champlain, [Vermont]). Sexual differences are visible in the shell; Alasmodon De Zool. New corrugata Kay 1843, females are slightly more swollen in the York, Moll., pt. 5: 198, pi. 24, fig. 259 (Passaic region of the posterior ridge. Occasionally River, New York; type, New York Lyceum of the is so that the lateral Nat. Hist, [destroyed by fire]). swelling great Mya rugulosa Wood 1856, in Hanley, Index faces of the shell in front of the ridge ap- 1 7 North Test, p. 199, pi. supp., fig. ( America; pear flat or even concave. The ventral type [probably lost]). margin be slightly emarginate, but Alasmidonta varicosa (Lamarck). Simpson, 1914, may shells Cat. Naiades, 1: 506. male are also sometimes emarginate Alasmidonta (Decurambis) varicosa (Lamarck). so that the sex can not alwavs be ascer- Ortmann, 1919, Mem. Carnegie Mus., 8: 190, tained from the shell alone. pi. 12, fig. 5. Clarke and Berg, 1959, Cornell Univ. Exp. Sta. Mem. no. 367, p. 28, fig. 34. Width Lengthmm Heightmm mm Athearn and Clarke, 1962, Natl. Mus. Canada,

Bull. 183, p. 25, pi. 3, figs. 5, 6. 73.0 40.0 31.0 N Fork, Shenandoah Biver, E Woodstock, Description, Shell usually small, from 40 Shenandoah Co., Vir- to 65 mm, though sometimes reaching 80 ginia. Female. mm in length. Outline rhomboid or sub- 44.0 26.0 18.0 Turkey Creek, 8 mi. NW Edgefield, Edgefield Co., elliptical. Valves not much inflated, thin South Carolina. Female. but strong. Anterior end regularly rounded; 33.0 28.0 12.0 Bocky Biver, 11 mi. N end and bi- posterior slightly elongated Sanford, Chatham Co., angulate. Ventral margin straight or North Carolina. Male. slightly arcuate. Dorsal margin straight, 32.0 29.5 13.0 As above. Female. forming an indistinct angle with the to Ortmann 1919: obliquely descending posterior margin. Anatomy. According ( 191 the of this is Hinge ligament located posteriorly of the ) , anatomy species very umbos, short but prominent. Posterior similar to that of Alasmidonta marginata ridge broadly rounded, with a second very Say. faint ridge above it, ending in a biangu- Breeding season. Ortmann (1919: 191) lation near the base. Posterior slope broadly found that in Pennsylvania breeding begins Atlantic Slope Unionidae • Johnson 355

lin August, when eggs are present; glochidia based. Ortmann (1919: 193) personally are found in September, and are carried collected A. varicosa in the upper Catawba over the winter and discharged the follow- River, North Carolina, and Clench and ing May. Okkelberg collected a single specimen in Habitat. Lives among rocks on gravel 1929 in a tributary of the Savannah River, substrates; also on sandy shoals, especially South Carolina. Seemingly rare below the in rapids and riffles of small rivers and Potomac River, Maryland, A. varicosa is creeks. relatively abundant on the Northern At- Remarks. Alasmidonta varicosa (La- lantic Slope. marck), having rudimentary pseudocar- Range. Atlantic Slope: tributary of the dinals, a lack of lateral teeth, and radial upper Savannah River system, South Caro- wrinkles on the posterior slope, is dis- lina, north to the lower St. Lawrence River tinguishable from the other Alasmidonta of system, Canada. the Atlantic Slope region. It is replaced in the upper Tennessee and Cumberland river Specimens Examined systems by A. ravenelina (Lea) which Savannah River System differs from varicosa by being more elongate, by having a more broadly bian- Savannah River Drainage. South Caro- gulate posterior ridge, and by usually not lina: Turkey Creek, 8 mi. NW Edgefield, having more than a trace of radial wrinkles Edgefield Co. (MZUM). on the posterior slope. Cooper-Santee River System Alasmidonta varicosa is the Atlantic re- of A. which is placement marginata Say Catawba River Drainage. North Caro- found in the Interior Basin throughout the lina: Catawba River, Bridgewater, Burke the upper Mississippi drainage, including Co. (Ortmann, 1919: 193). Ohio, Tennessee and Cumberland river systems. In the Northern Atlantic Slope Cape Fear River System region, marginata is found in the Susque- Cape Fear River Drainage. North Caro- hanna River, Pennsylvania, and the upper lina: Rocky River, 11 mi. N Sanford, Chat- St. Lawrence Canada. A. River, marginata ham Co. has a very sharp posterior ridge and a truncated which renders posterior slope, James River System the shell subtrapezoidal; also, the rays James River Virginia: Calf- tend to be broken into spots. A. varicosa Drainage. River 1846: 407 as A. has a gradually rounded, broadly biangu- pasture (Conrad, marginata Say). late posterior ridge, a gently rounded pos- terior slope, and a rhomboidal shape. The Occoquan Creek System rays are never broken into spots; the shell is not as heavy, nor does it grow as large as Oeeoquan Creek Drainage. Virginia: does marginata. Though marginata pre- Broad Run [Fairfax Co.], 3 mi. W fers larger rivers while varicosa prefers Manassas, Prince William Co. smaller streams, they are nevertheless Potomac River System sometimes found together where their but are ranges overlap, they easily sepa- Potomac River Drainage. Virginia: rated. N Fork, Shenandoah River, E Woodstock, Simpson (1914, 1: 506) correctly re- Shenandoah Co. S Fork, Shenandoah corded this species from as far south as River, Riverton, Warren Co. Maryland: South Carolina, but I was unable to find Potomac River, Hancock, Washington Co. the material on which his record was (Ortmann, 1919: 193). 356 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Genus Anodonta Lamarck Clarke, 1962, Natl. Mus. Canada Bull. no. 183, p. 27, pi. 2, figs. 5, 6. Subgenus Anodonta s.s. Lamarck Anodonta marginata Say 1817, Nicholson's En- Anodonta Lamarck Memoires tie la Soc. 1799, cyclopedia, 2 [no pagination], pi. 3, fig. 5 d'Hist. Nat. de Paris, p. 87. ( very common in our rivers [presumably near Linnaeus. Mono- Type species, Mytilus cygneus Philadelphia, Philadelphia Co., Pennsylvania, Placed on the Official List of Generic typic. and here so restricted]; type, ANSP [lost]). Names in in 94. Re- Zoology, 1926, Opinion Simpson, 1914, Cat. Naiades, 1: 388 [partim]. and Declarations confirmed, 1959, Opinions The measurements in the description, as tran- rendered Int. Zool. 20 by Comm. Nomen., scribed into millimeters, indicate the type to 561. (28): 303-10, Opinion have been 51 mm in length and 38 mm in Ann. 8: Ortmann, 1912, Carnegie Mus., 286, height. The figured shell was 69 mm in length Anodonta s. s. is found in partim. primarily and 36 mm in height. The first set of di- the Palearctic region of Europe and Asia and mensions are those of a cataracta from a creek like the monoecious and type species may be environment, while the second are of the have flat umbones. The several western elongated form which is sometimes found in American of the Pacific species region clearly ponds, and which becomes more common in to Anodonta s. s. belong the northern range of cataracta. Anodonta teres Conrad 1834, New Fresh Water Subgenus Pygonodon Crosse and Fischer Shells United States, p. 47, pi. 7, fig. 2 (Santee Canal Co.], South Carolina Pyganodon Crosse and Fischer 1894, in Fischer [Berkeley [type, ANSP lost]). Simpson, 1914, Cat. and Crosse, Mission Sci. an Mexique, pt. 7, Naiades, 1: 390. 2: 518. Species listed: A. glohosa Lea; A. Anodon excnrvata De Zool. of New tabascens Morelet; A. nopalatensis Sowerby; A. Kay 1843, 233 grijalvae Morelet; A. glauca Valenciennes. York, Moll., pt. 5, p. 202, pi. 17, fig. Introduced as a section. (Shaker Pond, Niskayuna, Albany Co., New York; Passaic River [Paterson, Passaic Co., New Type species, Anodonta globosa Lea. Subsequent New York State now in designation, Frierson, 1927, Check List North Jersey]; type, Colin., American Naiades, p. 9, teste Errata et Cor- USNM [not yet located]). rigenda. Raised to a subgenus. Anodonta virgulata Lea 1857, Proc. Acad. Nat. Sci. Phila., 9: 86 (Roanoke River, Weldon Frierson used (1927: 14-16) Pyganodon [Halifax Co.], North Carolina; Washington Co., as a subgenus to include most of the Georgia). Lea, 1862, Jour. Acad. Nat. Sci.

Anodonta of middle and eastern North Phila., ser. 2, 5: 213, pi". 33, fig. 282; figured America. holotype USNM 86593 from Roanoke River. Lea, 1863, Obs. Unio, 9: 35. It is that when the Interior suspected Anodonta lacnstris Lea 1857, Proc. Acad. Nat. Basin taxa included in this are subgenus Sci. Phila., 9: 84 (Crooked Lake and Little are revised, it will be found that there Lakes [Herkimer Co.], New York). Lea, 1860, about an equal number of species in the Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 363, Interior Basin and Atlantic Slope region. pi. 62, fig. 188; figured type USNM [not located]; though a slightly smaller specimen, USNM 86597 marked Mohawk, New York. Anodonta (Pyganodon) cataracta cataracta ex Lewis, is labeled holotype. Lea, 1860, Obs. Say Unio, 8: 45. Plate 14: 3,4 Anodonta liallenheckii Lea 1858, Proc. Acad. Nat. Plate 15: 1 Sci. Phila., 10: 138 (Uphaupee [Uphapee] Creek, Macon Co., Georgia [Alabama]). Lea, Anodonta fluviatilis Cmclin, of authors. See note 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: under Lampsilis ochracca Say, p. 388. 50, 32, fig. 112; figured holotype USNM Anodonta cataracta Say 1817, Nicholson's Encyclo- pi. 86428. Lea, 1859, Obs. Unio, 7: 50. Simpson, pedia, 2 [no pagination], pi. 3, fig. 2 (deep 1914, Cat. 1: 392. Clench and Turner, part of a mill dam [presumably near Philadel- Naiades, Bull. Florida State Mus., 1: 186, phia, Philadelphia Co., Pennsylvania, and here 1956, pi. I- so restricted]; type, ANSP [lost]). Simpson, 9, fig. 1914, Cat. Naiades, 1: 386. Ortmann, 1919, Anodonta gcsnerii Lea, 1858, Proc. Acad. Nat. Mem. Carnegie Mus., 8: 152, pi. 10, fig. 5, pi. Sci. Phila., 10: 139 (Uphaupee [Uphapee] 11, fig. 1. Clarke and Berg, 1959, Cornell Univ. Creek, Macon Co., Georgia [Alabama]). Lea, Exp. Sta. Mem. 367, p. 39, fig. 46. Athearn and 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: Atlantic Slope Unionidae • Johnson 357

231, pi. 31, fig. 109; figured holotype USNM Umbos slightly swollen and moderately Obs. 7: 49. S6427. Lea, 1859, Unio, convex, located in the anterior third or Anodonta dariensis Lea 1S58, Proc. Acad. Nat. fourth of the shell, their sculpture consist- Sci. Phila., 10: 139 (Hopeton, near Darien of five to seven [Mcintosh Co.]: Swift Creek, near Macon ing double-looped bars, [Bibb Co.]; Flint River; all Georgia). Lea, not appreciably lower in front or behind 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: the sinus, but of uniform elevation. Perios- 230, pi. 28, fig. 99; figured holotype USNM tracum generally smooth and shiny, slightly 86600 from Swift Creek. Lea, 1859, Obs. Unio, roughened toward the and on 7: 48. Simpson, 1914, Cat. Naiades, 1: 394. margins the Surface of the shell Anodonta williamsii Lea 1862, Proc. Acad. Nat. posterior slope. Sci. Phila., 14: 169 (Potomac River at the sometimes uniformly straw yellow, but Wbite House [District of Columbia]; below usually lighter to darker green with con- Mount Vernon Oth- [Fairfax Co., Virginia]; centric lighter and darker bands, with calooga Creek [Gordon Co.], Georgia. Lea, darker rays on the disk. The rays on the 1866, Jour. Acad. Nat. Sci. Phila., ser. 2, 6: are darker, at least those 27, pi. 10, fig. 26; type, USNM [not located]. posterior slope Lea, 1867, Obs. Unio, 11: 31. rays on the faint ridges, though generally Anodonta tryoni Lea 1862, Proc. Acad. Nat. Sci. the whole posterior slope is blackish green. 14: 169 River above Phila- Phila., (Schuylkill No hinge plate or teeth; muscle scars delphia; Delaware River at League Island inconspicuous and poorly defined. Nacre [both Philadelphia Co., Pennsylvania]; Flem- bluish white and iridescent. ington [ = Farmington, Hartford Co.], Con- necticut; Westfield [Hampden Co.], Massa- Females are sometimes indicated by a Potomac River near Chain chusetts; Bridge, swelling of the valves posterior to the above Washington, D. C). Lea, 1866, Jour. middle, but this is not a uniform character. Acad. Nat. Sci. Phila., ser. 2, 6: 28, pi. 10, fig. 27; type, USNM [not located], paratype, Length Height Width USNM 86572 from Potomac River. Lea, 1867, mm mm mm Obs. Unio, 11: 32. 165 86 62 Lake Michie, 12 mi. NNE Anodonta dolcaris Lea 1863, Proc. Acad. Nat. Durham, Durham Co., Sci. Phila., 15: 193 (Stewart's Mill Dam, North Carolina. Union Co., North Carolina). Changed to: 152 79 65 Savannah River, Johnsons Anodonta doliaris Lea 1866, Jour. Acad. Nat. Landing, 10 mi. W Allen- Sci. Phila., ser. 2, 6: 24, pi. 8, fig. 23; figured dale, Allendale Co., South

holotype ANSP 126522a. Lea, 1867, Obs. Carolina ( MZUM ) . Unio, 11: 28. Simpson, 1914, Cat. Naiades, 107 58 49 Swift Creek, near Macon 1: 393. [Bibb Co.], Georgia. Holo- type of A. dariensis- Lea. Description. Shell medium to large, reaching over 165 mm in length. Outline Anatomy. Thoroughly discussed by subelliptical, more or less elongate. Valves Simpson (G. B., 1884) and commented of young specimens not much inflated, upon by Reardon (1929: 8, pi. 3, figs. those of older ones quite inflated, thin, 1-10). The glochidia are hooked. sometimes very thin. Anterior end regularly Breeding season. From August to May rounded; posterior end rather pointed. (Ortmann, 1919: 154). Ventral margin straight, or slightly curved, Habitat. Lives in rivers and creeks in very occasionally slightly incurved. Dorsal soft mud and sand; also in sand bars, margin slightly curved, often forming a generally in slowly moving water; prefers winglike angle where it meets the obliquely ponds. descending posterior margin. The posterior Remarks. In the Atlantic Slope region margin joins the ventral margin in a broad Anodonta cataracta Say can be confused point, somewhat below the medial line. in the lower St. Lawrence drainage with Hinge ligament long and low. Posterior Anodonta cataracta fragilis Lamarck (see ridge indistinct. Posterior slope slightly Athearn and Clarke, 1962: 28; 1963: 22); concave, sometimes with two faint ridges. Anodonta implicata Say under which see: 358 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Remarks on p. 361; and in central New In the southern part of the Atlantic York with Anodonta grandis Say, of the Slope A. cataracta is not very common. In Interior Basin. A. cataracta Say differs the Altamaha River, at the stations listed from grandis primarily in the umbonal below, only a few specimens were taken. sculpture, that of the latter consists of four Lea's specimen of dariensis from Hopeton, to five bars of which the first two are near Darien, Mcintosh Co., Georgia, was concentric, while the others are distinctly not located, hence it is not included in the double-looped, with a sharp re-entering records since it might be gibbosa. In the sinus between the loops. The anterior loop Museum of Comparative Zoology there is is broadly rounded, the posterior loop is a single specimen of cataracta from the C. angular, narrow, and characteristically ele- B. Adams collection made before 1850, vated into a tubercle. In cataracta the bars labeled Washington Co., Georgia. If this are of uniform height and are not record is valid, then cataracta is found tubercular. These distinctions were prop- either in the upper Oconee River of the erly pointed out by Marshall (1890: 188, Altamaha River system or in the upper 189). Except for this minor difference the Ogeechee River system. This species be- two species are very close, though cataracta comes abundant and reaches large size in tends to be a brighter green, and while impounded waters, such as Lake Moultrie short high examples of cataracta are found of the Cooper-Santee River system, South in rivers, these forms are more common in Carolina, Stewarts Mill Dam, Union Co., grandis. In general, cataracta is more North Carolina, and Lake Michie on Flat elongated, especially individuals from River, North Carolina. ponds, and the posterior point is more Range. Alabama-Coosa River system. elevated above the base line, rendering Apalaehicolan region: Choctawhatchee and the ventral margin more convex. Ortmann upper Apalachicola River systems. Atlantic (1919: 154) has clearly pointed out the Slope: Altamaha River system, Georgia, different reaction of the two species to north to the lower St. Lawrence River environment. system, Canada. Interior Basin: probably Specimens from the Alabama-Coosa extending from the St. Lawrence River River system show no morphological dif- system westward to Michigan. ferences separating hallcnbeckii (Lea) (Clench and Turner, 1956: 186) from Specimens Examined cataracta. It is possible that cataracta ar- Altamaha River System rived there through the commingling of the upper Alabama and Coosa rivers di- Ocmulgee River Drainage. Georgia: rectly, though it is suggested elsewhere Possum Lake, NE Stone Mountain, De in this paper that cataracta may more Kalb Co. Swift Creek, near Macon, Bibb likely have spread into the Apalachicola Co. Limestone Creek, near Hartford, River system through a confluence of it Pulaski Co. (H. D. Athearn). Dicksons with the Savannah, and then subsequently Creek, 10 mi. NE Fitzgerald, Ben Hill Co. reached the Alabama system through a Ocmulgee River, below Lumber City, Tel- mingling of the waters of its Uphapee fair Co. (H. D. Athearn). Creek and Uchee Creek of the Apalachicola Savannah River System River system. Most of the shells from Uphapee Creek are the long pond forms. Savannah River Drainage. Sotitli Caro- I did not find any Anodonta in this broad, lina: Savannah River, 6 mi. S Martin; swift flowing creek in 1964, suggesting Savannah River, Johnsons Landing, 10 mi. that the original lot was from a quiet back- W Allendale; both (MZUM); both Allen- water which I did not locate. dale Co. Atlantic Slope Unionidae • Johnson 359

Cooper-Santee River System Rappahannock River System South Caro- Congaree River Drainage. Rapidan River Drainage. Virginia: lina: Congaree River. Mountain Run, Orange Co. (MZUM). Catawba River Drainage. North Caro- Potomac River System lina: Bissels Pond, Charlotte, Mecklenburg Co. (USNM). Potomac River Drainage. Pennsyl- Wateree River Drainage. Soutli Caro- vania: Great Tonoloway Creek, Thompson lina: Wateree River, 2.5 mi. W Camden, Township, Fulton Co. (Ortmann, 1919: Kershaw Co. 156). Conococheague Creek, Greencastle, Santee River Drainage. South Caro- Franklin Co. (Ortmann, 1919: 156). West lina: Poplar Creek, Schulers Fish Pond, Virginia: South Branch, Potomac River, near Santee State Park; Eutavv Springs, Romney, Hampshire Co. (Ortmann, 1919: [town of] Eutaw Springs; both Orange- 157). Virginia: South River (upper Shen- burg Co. Lake Moultrie, Cross, Berkeley andoah River), Waynesboro, Augusta Co. Co. (Ortmann, 1919: 157). Potomac River, Great Falls Co. Fairfax Co. District Pedee River System Park, of Columbia: Aqueduct Lake. Yadkin River Drainage. North Caro- lina: Co. Yadkin Salem, Forsyth (USNM). Anodonta (Pyganodon) gibbosa Say River, Rowan Co. Stewarts Mill Dam, Plate 15:2,3 Union Co. (USNM). Anodonta gibbosa Say 1824 in W. H. Keating, Waccamaw River System Narrative of an Expedition to the Source of the St. Peters River, 2: 265, pi. 14, figs. 3-4 Waccamaw River Drainage. North (South Carolina; type, ANSP [lost]. Type Carolina: Lake Waccamaw, [town of] locality is here restricted to Altamaha River, Lake Waccamaw, Columbus Co. Hopeton, near Darien, Mcintosh Co., Georgia. See: Remarks helow). Simpson, 1914, Cat. Cape Fear River System Naiades, 1: 397. Anodonta gibbosa Say, partim. Clench and Fear River North Caro- Cape Drainage. Turner, 1956, Bull. Florida State Mus., 1: 184. lina: Travis Creek, 1.5 mi. N Gibsonville, Description. Shell medium to large, Guilford Co. (MZUM). Rocky River, 11 reaching 120 mm in length. Outline sub- mi. N Sanford, Chatham Co. Livingstons elliptical. Valves very much inflated, thin Creek [Brunswick Co.] (USNM). and smooth. Anterior end regularly Neuse River System rounded; posterior end broadly rounded or somewhat pointed. Ventral margin usually Neuse River Drainage. North Carolina: broadly curved, though centrally some- Lake Michie, 12 mi. NNE Durham Center, times straight, or slightly incurved. Dorsal Durham Co. (75). margin slightly curved, usually forming a Roanoke River System distinct winglike angle where it meets the margin. The Dan River Drainage. Virginia: Aarons obliquely descending posterior the ventral margin Creek, 3 mi. W Buffalo, Lithia Springs; posterior margin joins in a broad near the medial line. Dan River; both Halifax Co. point and low. Posterior Roanoke River Drainage. North Caro- Hinge ligament long rounded, lina: Roanoke River, Weldon, Halifax ridge broadly though occasionally with an Posterior Co. (USNM). a trace of angle. slope slightly concave. Umbos very high and James River System broad, located in the anterior third of the James River Drainage. Virginia: Calf- shell, their sculpture consisting of a num- pasture River (Conrad, 1846: 407). ber of low double-looped ridges. Perios- 360 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

tracum generally smooth and shiny, slightly a specimen in the Museum of Comparative roughened toward the margins and on Zoology from B. Walker, collected at Hope- ton the lower has almost the the posterior slope. Surface of the shell on Altamaha, the To avoid straw yellow to greenish, older specimens same measurements as type. becoming chestnut to blackish. Usually confusion the restriction of the type lo- to this seems warranted. with very fine greenish rays; the several cality region Turner included on the posterior slope usually darker and Clench and (1956: 185) broader than those on the disk. under this name the Anodonta of the and Chocta- No hinge plate or teeth; muscle scars Ochlockonee, Apalachicola inconspicuous and poorly defined. Nacre whatchee River systems, but these Ano- bluish white and iridescent, sometimes donta are more elliptical and elongate, less with the more pinkish toward the umbonal cavities. much globose, and tuberculous beak sculpture of Anodonta Length Height Width mm mm mm grandis Say, which they appear to be. 123 72 69 Altamaha River, 3 mi. NW Range. Southern Atlantic Slope: re- Everett City, Glynn Co., stricted to the Altamaha River system, Georgia. Georgia. 97 59 48 Cedar Creek, Fountains Mill, 7 mi. SW Hawkins- Specimens Examined ville, Pulaski Co., Georgia. 74 49 40 [after Say]. Altamaha River System 71 47 37 Altamaha River, Hopeton, River Drainage. Georgia: near Darien, Mcintosh Co., Ocmulgee Cedar Fountains 7 mi. SW Georgia. Creek, Mill, Hawkinsville, Pulaski Co. Ocmulgee River, Habitat. Lives in soft mud and sand, 1 mi. S Lumber City, Telfair Co. also in sand in mov- bars; generally slowly Altamaha River Drainage. Georgia: water. ing Altamaha River, 10 mi. NE Surrency, Ap- Remarks. In the Atlantic Slope region, pling Co. Altamaha River, "Riverside Park," Anodonta gibbosa Say can only be con- 4 mi. N Jesup, Wayne Co. Altamaha River, fused with A. cataracta Say, but gibbosa 3 mi. NW Everett City, Glynn Co. Button- has a swollen the umbos are very shell, wood Swamp, Liberty [now Long] Co. and the swelling con- greatly protruded, Altamaha River, Hopeton, near Darien, tinues with the of the the growth shell, Mcintosh Co. greatest swelling extending toward the ventral margin. Frierson (1912: posterior Anodonta {Pyganodon) implicata Say out the 129) correctly pointed relationship Plate 15: 4 of this to A. cataracta based species Say Plate 16: 1,2 on the beak sculpture, but thought gibbosa Anodonta implicata Say 1829, New Harmony might be merely a subspecies. They ap- [Indiana] Disseminator 2, no. 22, p. 340 (pond to be and both occur pear separate species in Danvers [Essex Co.], Massachusetts; type, in the Altamaha River system. ANSP [lost]). was to It is obvious that Major Le Conte who The type locality changed Agawani River (outlet of Halfway Pond), Plymouth sent T. Say the type of gibbosa made a [Plymouth Co.], Massachusetts, and a neotype mistake as to its since there are locality, MCZ 176769 was selected by Johnson, 1946, no of this in of the specimens species any Occ. Papers on Moll., 1: 112, pi. 16, figs. 1, 2). collections studied from South Carolina. Simpson, 1914, Cat. Naiades, 1: 391. Ortmann, 8: Le Conte was to later send I. Lea numerous 1919, Mem. Carnegie Mus., 159, pi. 11, figs. 2, 3. Clarke and Berg, 1959, Cornell species he collected in the lower Altamaha Univ. Exp. Sta. Mem., no. 367, p. 40, fig. 42. so it is that this River, presumed species Athearn and Clarke, 1962, Natl. Mus. Canada, was also from this region, especially since Bull. no. 183, p. 26, pi. 2, figs. 1, 2. Atlantic Slope Unionidae • Johnson 361

Anodonta newtonensis Lea Trans. Amer. 1838, anterior margin below the pallial line. Philos. Soc, 6: 79, pi. 21, fig. 66 (Newtown Nacre dull opalescent, generally pale cop- [Newton] Creek, [Camden Co., New Jersey], per, pinkish, or, more white or opposite Philadelphia; figured type not in rarely, bluish with a bluish cast USNM or ANSP [lost]; Schuylkill [River], white, sometimes Fairmonnt [Philadelphia, Philadelphia Co., toward the margins. USNM 86561 selected Pennsylvania], lectotype Females are usually more swollen than by Johnson, 1946, Oec. Papers on Moll., 1: 112. males in the middle portion of the disk, [The selection of lectotypes, without figuring which tends to make the lower them, is a dubious contribution to knowledge, margin more curved the shell if, indeed, such selections are valid at all.] and somewhat Figured here. Pi. 15: 4. Lea, 1838, Obs. Unio, shorter and higher than that of the male, 2: 79. but this is not a uniform character. Anodonta housatonica Linsley 1845, Amer. Jour. ser. 48: 277 Width Sci., 1, (Housatonic [River], Lengthmm Heightmm mm Corum [near Huntington, Fairfield Co.], Con- necticut), [nomen nudum]. Gould, 1848, Amer. 162 77 66 Agawam River, Plymouth, Jour. Sci., ser. 2, 6: 234, figs. 4, 5; measured Plymouth Co., Massachu- and figured holotype USNM 678302. setts. Male. 120 60 48 As above. Neotype of A. Shell medium to Description. large, implicata Say. Male. 165 mm in Outline elon- reaching length. 97 52 39 Schuylkill River, Fair- gate-elliptical to elongate-ovate. Valves mount, Philadelphia Co., quite inflated, somewhat subcylindrical, Pennsylvania. Lectotype of A. newtonensis Lea. Fe- rather solid and heavy for an Anodonta. male. Anterior end regularly rounded; posterior 94 51 34 Housatonic River, Corum, end more older acutely rounded, speci- Fairfield Co., Connecticut. mens often becoming subtruncated. Ven- Holotype of A. housatonica tral margin slightly rounded, becoming Gould. Male. straight or slightly arcuate in old individu- Anatomy. The anatomy of this species als. Dorsal margin straight, occasionally has not been investigated. The glochidia forming a slight winglike angle where it are rather large, subtriangular, with a meets the obliquely descending posterior spine at the tip of each valve. The host margin. Hinge ligament long and low. fish is the alewife, Alosa pseudoha renins Posterior ridge not very acute, but gener- (Wilson) (Johnson, 1946: 112). ally distinctly biangulate. Posterior slope Breeding season. According to Ortmann slightly concave. Umbos slightly swollen (1911: 303) all Anodonta are bradytictic, and moderately convex, located in the an- or winter breeders, the marsupia filling terior third of the shell, their sculpture sometime in summer between July and of five to seven consisting double-looped September. The glochidia are carried not lower in or bars, appreciably front through the winter and set free when behind the sinus but of uniform elevation. warmer weather begins. I collected gravid Periostracum generally rather smooth, save females at Plymouth, Massachusetts, on for lines growth and occasional plaiting, May 8, 1943, and June 9, 1946. yellowish brown or greenish brown, some- Habitat. Lives in sand or gravel, rarely times becoming reddish brown or almost in mud. Found abundantly, close to the black in old specimens. Immatures are coast in ponds that have an unobstructed sometimes greenish and obscurely rayed. outlet to the ocean. Seems to prefer a No hinge plate or teeth. Beak cavities stream environment; the largest examples rather shallow. Muscle scars and pallial have been taken in a relatively swift, sandy line distinct especially in mature speci- stream. mens. Shell distinctly thickened along the Remarks. Anodonta implicata Say of the 362 Bulletin Museum of Comparative Zoology, Vol 140, No. 6

Northern Atlantic Slope can generally be mann and Walker, 1922: 32), Frierson included all of the taxa mentioned in the easily separated from all other species of Anodonta. The most reliable and constant present paper, except A. suborbiculata Say which he intro- feature in identifying adults of this species of the Interior Basin for is the pronounced thickening of the an- duced the subgeneric name Utterbackiana, terior-ventral portion of the shell below on the basis that that species is dioecious. is al- as mentioned this as an un- the pallial line. This thickening not But, below, ways pronounced in immatures, but may reliable basis for classification and the still be detected. Other characteristics of shell morphology is clearly that of Utter- this species are the dark and usually yel- backia. 1956: states lowish or brownish rayless periostracum Morrison (in Walter, 265) like A. of and the pale copper or salmon color of the that Anodonta imbecilis, cygnea nacre. Europe and Asia, is monoecious and has that therefore imbecilis Anodonta implicata is especially abun- flat umbos, and dant in the region of Cape Cod, Massachu- belongs to Anodonta s. s. that setts, where it reaches its greatest size. The Heard (1966: 31) has clearly shown tide waters of the Potomac River, Virginia, sexuality is an unreliable means of classi- seem to be its southern terminus. All the fying Anodonta. Neither cygnaea nor im- specimens under this name I have seen becilis are uniformly monoecious. The flat from rivers below the Potomac were heavy umbos of Utterbackia and Anodonta s. s. cataracta, which have an especially distinct appear to be a convergent character. Utter- is isolated s. s. pallial line anteriorly, but which lack the backia quite from Anodonta restricted special anterior-ventral thickening of the in North America, the latter being shell and the characteristic nacre color. It to the Pacific region. All four species of Ut- is on the basis of such specimens that terbackia are more delicate than cygnea, and Atheam and Clarke (1962: 26) suggest some individuals of each of the species that this species extends to South Carolina. exhibit fine rays toward the umbos. These Range. Northern Atlantic Slope: Potomac are lacking in cygnea or any other Ano- River, Maryland, north to New Brunswick donta. and northern Nova Scotia, both Canada. There are two species of Utterbackia in the Interior Basin, suborbicidata and im- Specimens Examined becilis. The latter is found almost every- where to the east Peninsular Florida Potomac River System except and the Northern Atlantic Slope region. Potomac River Drainage. District of Speciation has taken place in the South- Columbia: Potomac River (USNM). eastern states, where two species, in addition to imbecilis, occur. They are F. Baker Subgenus Utterbackia C. couperiana Lea, covered here, and A. Utterbackia F. C. Baker American Midland 1965 which occurs in 1927, peggyae Johnson ( ) , Nat., 10: 221, 222 (misspelled as Utterbackia the Apalachicolan region and Gulf drain- on p. 221). age of Peninsular Florida. Type species, Anodonta imbecillis [sic] Say. Orig- inal designation. imbecilis Utterbackiana Frierson 1927, Check List North Anodonta (Utterbackia) Say American Naiades, p. 17. Plate 16: 3 Type species, Anodonta suborbiculata Say. Mono- Anodonta (Lastena) ohiensis Rafinesque 1S20, typic. Ann. Gen. Sci. Physiques (Bruxelles), 5: 316 et toutes les rivieres Under the subgenus Lastena Rafinesque (l'Ohio adjacentes; sup- posed type, ANSP 216908, is Proptera laevissima available for use here since the (not type (Lea)). Ortmann and Walker (1922, Occ. is Anodonta lata Rafinesque, teste Ort- Papers, Mus. Zool. Univ. Michigan, no. 112, p. Atlantic Slope Unionidae • Johnson 363

A. is not identi- 37) clearly showed that ohiensis margin slightly convex, often almost straight nevertheless Frierson ( 1927, Check List fiable, in the middle. Dorsal margin straight and North American Naiades, p. 18) persisted in long, usually forming a distinct angle using this name For A. imbecilis Say. where it the Anodonta imbecilis Say 1829, New Harmony meets obliquely descending [Indiana] Disseminator, 2(23): 355. (Wabash posterior margin. The posterior margin ANSP River, Indiana; type, [lost]). Neotype joins the curved ventral margin in a point Senckenberg Mus. 4301 [not seen] selected, near the medial line. Hinge ligament short but not figured, by Haas, 1930, Senckenberg- and inconspicuous. Posterior iana, 12: 326. Type locality restricted to ridge broadly Wabash River, New Harmony, Indiana, by rounded, though occasionally rather an- Clench and Turner, 1956, Bull. Florida State gular. Posterior slope slightly concave. Mus., 1: 188. Umbos low and broad, seldom extending Anodonta incerta Lea 1834, Trans. Amer. Philos. above the dorsal margin, located in the Soc, 5: 56, pi. 6, fig. 16 (Ohio River, near anterior third of the their Cincinnati [Hamilton Co., Ohio]; type, not in shell, sculp- USNM or ANSP [lost]. Lea, 1834, Obs. Unio, ture consisting of a number of delicate 1: 158. subconcentric undulations. Periostracum Anodon horda Could 1855, Proc. Boston Soc. Nat. smooth and shiny, except the posterior Hist., 5: 229 (Comanche Creek [Tributary of slope which be slightly roughened. The Llano River, close to the present site of may Surface of the shell straw to Mason, Mason Co.], Texas [teste, Taylor, 1967, yellow green- Veliger, 10: 153]; measured holotype USNM ish yellow, but more generally darker 678301). green, sometimes grass green. Toward the Anodonta henryana Lea 1857, Proc. Acad. Nat. umbos the color may be greyish, brownish Sci. Phila., 9: 102 (Matamoras, Tamaulipas or with concentric darker [State], Mexico). Lea, 1860, Tour. Acad. Nat. yellowish, disk be Sci. Phila., ser. 2, 4: 373, pi. 66, fig. 198; growth-rests. The may obscurely figured holotype USNM 86692. Lea, 1860, rayed; the posterior slope usually has two Obs. 8: 55. Unio, or three dark green to blackish rays. Utterbackia imbeciUis [sic] fusca Baker 1927, No hinge plate or teeth; muscle scars American Midland Nat., 10: 222 (Sturgeon and defined. Nacre Bay, Door Co., Wisconsin; holotype MZUM inconspicuous poorly 209141, figured by Baker, 1928, Bull. Wisconsin bluish white and iridescent, sometimes Geol. Nat. Hist. Survey, no. 70(2): 175, pi. 103, pinkish toward the umbonal cavities. 14). fig. This species is hermaphroditic and does Anodonta imbeoillis 1914, [sic] Say. Simpson, not show anv sexual differences. Cat. Naiades, 1: 396. Anodonta henryana Lea. Simpson, 1914, Cat. Length Height Width Naiades, 1: 396. mm mm mm Anodonta ohiensis Rafinesque. Ortmann, 1919, 90 42 34 Magnolia Springs, Mag- Mem. Carnegie Mus., 8: 162. nolia State Park, Perkins, Anodonta imbecilis Say, partim. Clench and Jenkins Co., Georgia. Turner, 1956, Bull. Florida State Mus., 1: p. 84 44 31 Outlet of Waynesboro 187. See: lohnson, 1965, Breviora, Mus. Comp. Lake, Waynesboro, Burke Zool., no. 213, pi. 2, fig. 5, 6. Co., Georgia. Anodonta imbecilis Clarke (Utterbackia) Say. 71 39 24 Buck Creek, 5 mi. NW and 1959, Cornell Univ. Sta. Mem. Berg, Exp. Hawkinsville, Pulaski Co., no. 42. 367, p. 40, fig. Georgia. Shell small to Description. medium, Anatomy. Clarke (1959: 42) discusses usually not reaching over 90 mm in length. the work that has been done on the anat- Outline rather elongated, subelliptical. omy of this species. He suggests that the Valves compressed, or only slightly swollen host fish is Semotilus atromaculatus atro- in young shells, but often becoming quite maculatus (Mitchill). inflated with age, very thin, fragile, and Breeding season. Ortmann (1919: 163) smooth. Anterior end regularly rounded; says this species is undoubtedly brady- posterior end somewhat pointed. Ventral tictic. 364 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Habitat. Lives in soft mud or sand in Northern Atlantic Slope: Gunpowder River ponds, creeks, and near the banks of larger system, Maryland. rivers. Remarks. In the Atlantic Slope region, Specimens Examined be con- Anodonta imbedlis Say can only Altamaha River System fused with A. coupcriana Lea and in the River Apalachicolan region it can only be mis- Ocmulgee Drainage. Georgia: Buck 5 mi. NW Hawkinsville; taken for Anodonta peggyae Johnson. Creek, Cedar Creek, Fountains Mill, 7 mi. WSW These three species of Anodonta are the Hawkinsville; both Pulaski Co. only ones in these drainage systems whose umbos do not extend above the dorsal Ohoopee River Drainage. Georgia: 3 mi. margin. In imbedlis the ventral margin is Yam Grande Creek, W Swainsboro, Emanuel Co. almost parallel to the dorsal one, while in River couperiana the ventral margin is broadly Altamaha Drainage. Georgia: curved, which renders the shell much Altamaha River, 4 mi. NE Jesup, Wayne Co. higher in proportion to its length than imbecilis. See: Remarks under Anodonta Ogeechee River System couperiana Lea on p. 365. The general distribution of A. imbecilis Ogeeehee River Drainage. Georgia: has been discussed by Ortmann (1919: Magnolia Springs, Perkins, Jenkins Co. 165). It is found in the Apalachicolan Mill Creek, 5 mi. E Midville; Barkcamp region east to the Ochlockonee River Creek, 7 mi. E Midville; both Burke Co. system, Florida. In the Atlantic Slope Savannah River System region it does not extend below the Alta- maha River system, Georgia. It apparently Savannah River Drainage. Georgia: does not occur in Peninsular Florida, since Savannah River, 7.5 mi. NE Shellbluff; the author made an extensive collecting outlet of Waynesboro Lake, Waynesboro; to this area in did find trip 1962 and not Savannah River, 6 mi. NE Girard; all A. imbecilis. It is on the Gulf replaced Burke Co. South Carolina: Savannah River, side of Florida Anodonta. by peggyae John- 6 mi. S Martin; Savannah River, Johnsons son (1965: 1) and by A. couperiana Lea on Landing, 10 mi. W Allendale; Savannah the Atlantic side. (See: Johnson, 1965, River, Kingjaw Point, 10 mi. WSW Allen- pi. 1.) On the Atlantic slope, A. imbecilis dale (ANSP); all Allendale Co. extends to the Nottoway River, Virginia, of the Chowan River system. Ortmann Cooper-Santee River System (1919: 163) records this species from Catawba River North Caro- Pennsylvania in the region of Lake Erie, Drainage. lina: Bissels Pond, Charlotte, but not from the Atlantic Slope. It has Mecklenburg Co. since been found at Gunpowder River, (ANSP; USNM). Loch Raven Dam, Loch Raven, Baltimore Wateree River Drainage. South Caro- Co., Maryland (ANSP). lina: Wateree River, 2.5 mi. W Camden, Range. Interior Basin: Mississippi drain- Kershaw Co. age generally, Ohio River, West Gulf Santee River Drainage. South Carolina: Coastal region, Alabama-Coosa River Santee River, above dam, near Cross; system; and Apalachicolan region: Rio Charleston (USNM); both Berkeley Co. Grande River system, Texas, east to the Pedee River System Ochlockonee River system, Florida. South- ern Atlantic Slope: Altamaha River system, Yadkin River Drainage. North Caro- Georgia, north to the Chowan River system. lina: Salem, Forsyth Co. (USNM). Atlantic Slope Unioniime • Johnson 365

Cape Fear River System rounded. Posterior slope slightly concave. Umbos low and broad, seldom extending Cape Fear River Drainage. North Caro- above the dorsal margin, located in the lina: Greenfield Mill Pond, Wilmington, anterior third of the shell, their sculpture New Hanover Co. (Ortmann, 1919: 164). consisting of a number of delicate subcon- Chowan River System centric undulations. Periostraeum smooth and shiny, except the posterior slope which Nottoway River Drainage. Virginia: may be roughened. Surface of the shell Birchen Lake, on Birchen Creek, Camp straw yellow to yellowish green, usually Pickett, Nottoway Co. with numerous and generally fine green Gunpowder River System rays, sometimes with distinctly darker rays on the posterior slope. Gunpowder River Drainage. Maryland: No hinge plate or teeth; muscle scars Gunpowder River, Loch Raven Dam, Loch inconspicuous and poorly defined. Nacre Baltimore Co. Raven, (ANSP). bluish white and iridescent.

Width Anodonta {Utterbackia) couperiana Lea Lengthmm Heightmm mm Plate 16:4 110 67 49 Sandpit, near Edisto Biver, Plate 17: 1 Givhans Ferry State Park, 2 mi. NW Givhans, Dor- Anodonta cowperiana [sic] Lea 1840, Proc. Amer. chester Co., South Caro- Philos. Soc, 1: 289 (Hopeton, near Darien lina. [Mcintosh Co.], Georgia). Changed to: 35 near Anodonta couperiana Lea 1842, Trans. Amer. 87 50 Pond, Savannah, Chatham Co., Georgia. Philos. Soc., 8: 227, pi. 20, fig. 46; figured type, not in USNM [lost]. Lectotype, USNM 83 46 35 Charleston, South Carolina. 86673, selected by Johnson, 1965, Breviora, Lectotype of A. dunlapiana Mus. Comp. Zool., no. 213, p. 3, pi. 2, fig. Lea. 4. Lea, 1842, Obs. Unio, 3: 65. 67 38 29 Hopeton, near Darien, Mc- Anodonta dunlapiana Lea 1842, Proc. Amer. intosh Co., Georgia. Lecto- Philos. Soc, 2: 225 (South Carolina). Lea, type of A. couperiana Lea. 1842, Trans. Amer. Philos. Soc., 8: 248, pi. Discussed Lea 27, fig. 65; figured type, not in USNM [lost]. Anatomy. by (1863: 451). Lectotype, here selected, USNM 86564, Pi. 17, Habitat. Prefers sandy or muddy bot- Chatham Caro- fig. 1, [Charleston, Co.] South toms of ponds and sluggish streams. lina. Lea, 1842, Obs. 3: 86. Unio, Remarks: In the Southern Atlantic Slope Anodonta cowperiana [sic] Lea. Clench and region, Anodonta couperiana Lea can be Turner, 1956, Bull. Florida State Mus., 1: 183, confused only with A. imbecilis Say. Both pi. 6, fig. 3. species have umbos which do not extend Description. Shell medium to large, above the dorsal margin, a character which reaching 110 mm in length. Outline sub- distinguishes them from the other Ano- elliptieal to subcircular. Valves somewhat donta of the Atlantic Slope. A. couperiana inflated, thin, fragile, and smooth. Anterior is differentiated from imbecilis by its green end regularly rounded; posterior end rays, which are especially fine on the disk, somewhat pointed. Ventral margin broadly- and by its broadly curved ventral margin, curved. Dorsal margin straight and long, which renders the shell much higher in usually forming a distinct winglike angle proportion to its length than imbecilis, where it meets the obliquely descending whose ventral margin is almost straight posterior margin. Hinge ligament short but and parallel to the dorsal one. The prominent. The posterior margin joins the height/length ratio of couperiana is about curved ventral margin in a point near the 2 to that of 1.5 in imbecilis. medial line. Posterior ridge broadly A. couperiana is found in the Apalach- 366 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

icola River system with A. peggyae John- Edisto River System which differs from son, couperiana by being Edisto River Drainage. South Caro- subrhomboidal rather than in elliptical lina: Sandpit, near Edisto River, Givhans and more outline, by being biangulate pos- Ferry State Park, 2 mi. NW Givhans, with the near teriorly, biangulation ending Dorchester Co. Charleston, Chatham Co. is more the base. A. couperiana pointed (USNM). posteriorly with the point ending near the medial line. In A. peggyae, when the dorsal Cooper-Santee River System margin is held straight, the ventral margin Saluda River Drainage. South Carolina: is often toward the obliquely descending Milton, Laurens Co. broad posterior basal biangulation. A. couperiana is a rare species in Geor- Cape Fear River System gia, but the specimens from the northern Cape Fear River Drainage. North Caro- part of the range are remarkable for their lina: Greenfield Mill Pond, Wilmington, size and It is common and abun- beauty. New Hanover Co. (USNM). dant in Central Florida, where individuals tend to be small. Genus Strophitus Rafinesque Range. Apalachicolan region: Apalach- Strophitus Rafinesque 1820, Ann. Gen. ties Sei. icola, Ochlockonee, and St. Marys river Physiques (Bruxelles), 5: 316. Peninsular Florida. Southern At- systems. Type species, Anodonta undulata Say. Monotypic. lantic Slope: Altamaha River system, Geor- Ortmann, 1912, Ann. Carnegie Mus., 8: 299. Frierson Cheek List North gia, north to the Cape Fear River system, Pscudodontoideus 1927, North Carolina. American Naiades, p. 9, 23. Instituted as a subgenus. Type species, Margaritana alabamensis Lea. Specimens Examined Original designation.

Saint Marys River System Strophitus contains two species, undu- latus which is widely distributed Saint Marvs River Drainage. Florida: (Say), the Interior Rasin and Atlantic St. Marys River (ANSP; MZUM). throughout Slope region, and suhvexus (Conrad) Altamaha River System (Johnson, 1967: 3, pi. 1, figs. 1-3), which extends from the eastern part of the West Altamaha River Drainage. Georgia: Gulf Coastal region through the Apalach- Rice field ditches on the Altamaha River, icolan region. All of the taxa included Hopeton, near Darien, Mcintosh Co. by Frierson under Strophitus s. s. and Vseudodontoideus Frierson are included in Savannah River System the synonymy of the two species mentioned Savannah River Drainage. South Caro- above, except for: Strophitus undulatus lina: Brickyard Pond, Hamburg, Aiken Co. tennesscensis Frierson, which may be a (USNM). Savannah River [Johnsons geographic subspecies in the modern sense; Landing], 10 mi. W Allendale; Savannah and Strophitus radiatus (Conrad), which River, Kingjaw Point, 10 mi. WSW Allen- belongs under Anodontoidcs radiatus dale (ANSP); both Allendale Co. Georgia: (Conrad) (Johnson, 1967: 6, pi. 2, figs. Pond near Savannah, Chatham Co. 1-4). Generally, Frierson gave neither de- Combahee River System scriptions nor reasons for instituting new Salkehatchie River Drainage. South subgenera, but it is presumably on the Carolina: Pauline Cave, near Kline, Barn- basis of the vestigial pseudocardinal teeth in = that well Co. (ANSP). found alabamensis ( subvexus) Atlantic Slope Uniontoae • Johnson 367

Anodonta Lea 1852, Trans. Amer. he instigated Pseudodontoideus as a sub- sliacffcriana Philos. Soc, 10: 288, pi. 26, fig. 50 (Horn genus. If the present synonymy of the Lake Creek [Shelby Co.], Tennessee; figured is correct, does not need species Strophitus holotype USNM 86685). Lea, 1852, Obs. subdivision. Unio, 5: 44. Alasmodon rhombica Anthony 1865, Amer. Jour. 1: 5 Strophitus undulatus (Say) Conch., 158, pi. 12, fig. (Michigan; figured holotype MCZ 50296). Plate 17: 2, 3 Anodon pa))ijracca Anthony 1865, Amer. Jour. Anodonta undulata Say 1817, Nicholson's Encyclo- Conch., 1: 161, pi. 15, fig. 2 (locality unknown pedia, 2 [no pagination], pi. 3, fig. 6 (no type [Potomac River, Virginia]; figured holotype locality, here restricted: Schuylkill River, near MCZ 150656). Philadelphia, Philadelphia Co., Pennsylvania; Anodon annulatus Sowerby 1867, Conch. Iconica, type, AXSP [lost]). 17, Anodon, pi. 18, fig. 67 (Hab. ?; Cuming Anodonta pensylvanica [sic] Lamarck 1819, Hist. collection in BMNH [lost]). Nat. des Animaux sans Vertebres, 6: 86 (la Anodon quadriplicatus Sowerby 1867, Conch. riviere de Schuglkill [Schuylkill], pres de Phila- Iconica 17, Anodon, pi. 28, fig. 110 (Potomac delphie [Philadelphia Co., Pennsylvania]; holo- River, [Virginia]; holotype in Walpole coll., type, Geneva Museum, figured by Delessert, not in BMNH, [probably lost]). 1841, Rec. Coquilles de Lamarck, pi. 13, figs. Anodonta sahnonia Clessin 1873, Conch. Cab., 4a, 4b). ser. 2, 9, pt. 1: 91, pi. 24, figs. 1, 2 (Norda- Zool. Jion Lea Anodon wgosus Swainson 1822, Illustrations, merika, im Ohio-und Wabash-Fluss ) ; ser. 1, 2, pi. 96 (United States; type [location 1838.

unknown] ) . Strophitus undulatus ovatus Frierson 1927, Check Anodon areolatus Swainson 1829, Zool. Illustra- List North American Naiades, p. 22 (Middle tions, ser. 2, 1, pi. 18 (North America; type West [Lyon Creek, Edwards Co., Illinois]; syn- [location unknown]). type MZUM 87584). Alasmondonta edcntula Say 1829, New Harmony Strophitus rugosus pepinensis Baker 1928, Bull. [Indiana] Disseminator 2, no. 22, p. 340 Wisconsin Geol. Nat. Hist. Survey, no. 70(2), (Wabash River [near New Harmony, Posey Co., p. 204, pi. 74, fig. 8 (Lake Pepin near Lake Indiana]; type, ANSP [lost]). City [Wabasha Co.], Minnesota; holotype Anodonta virgata Conrad 1836, Monography MZUM 209137).

Unionidae, no. 5, back cover ( Buck Creek, Strophitus rugosus ivinnchagoensis Baker 1928, Clarke Co., Ohio; type ANSP [lost]). Bull. Wisconsin Geol. Nat. Hist. Survey, no. Anodonta pavonia Lea 1836, Trans. Amer. Philos. 70(2), p. 205, pi. 74, figs. 1-6 (Lake Winne- Soc, 6: 78, pi. 21, fig. 65 (headwaters of the bago, Long Point Island, Wisconsin; holotype Little Beaver [River, Lawrence Co.], Ohio; MZUM [original no. Univ. of Wisconsin 943a]). figured holotype USNM 86514). Lea, 1838, Stro))hitus rugosus lacustris Baker 1928, Bull. Obs. Unio, 2: 78. Wisconsin Geol. Nat. Hist. Survey, no. 70(2), Anodonta wardiana Lea 1838, Trans. Amer. Philos. p. 207, pi. 75, figs. 6-8 (Oconomowoc Lake, Soc, 6: 46, pi. 14, fig. 42 ([Scioto River], Waukesha Co., Wisconsin; holotype Univ. near Chillieothe [Ross Co.], Ohio; figured holo- Illinois, Urbana Z-22073 [not seen]). type USNM 86488). Lea, 1838, Obs. Unio, Strophitus edentulus (Say). Simpson, 1914, Cat. 2: 46. Naiades, 1: 345. Ortmann, 1919, Mem. Car- Anodon unadilla De Kay 1843, Zool. New York, negie Mus., 8: 197, pi. 12, figs. 7, 8. Moll., pt. 5: 199, pi. 15, fig. 228 (Unadilla Strophitus undulatus (Say). Simpson, 1914, Cat. River, a tributary of the Susquehanna [River Naiades, 1: 349. Ortmann, 1919, Mem. Car- system], Otsego Co., New York; type, New York negie Mus., 8: 195, pi. 12, fig. 6. Clarke and Lyceum of Nat. Hist, [destroyed by fire]). Berg, 1959, Cornell Univ. Exp. Sta. Mem., no. Anodonta tetragona Lea 1845, Proc. Amer. Philos. 367, p. 43, fig. 41. Athearn and Clarke, 1962, Soc, 4: 165 (Alexandria [Rapides Parish], Natl. Mus. Canada, Bull. no. 183, p. 29, pi. Louisiana). Lea, 1848, Trans. Amer. Philos. 3, figs. 7, 8. Soc, 10: 82, pi. 8, fig. 25; figured holotype seldom ex- USNM 86682. Lea, 1848, Obs. Unio, 4: 56. Descri))tion. Shell medium, Anodonta arkansensis Lea 1852, Trans. Amer. ceeding 90 mm in length in the Atlantic Philos. Soc, 10: 293, pi. 29, fig. 56 (Little boid to subrhomboid. Valves subinflated, Arkansas River, where the road to Santa Fe generally rather thin, inequilateral. An- crosses it [Sedgwick Co., Kansas]; figured holo- terior end rounded; posterior end type USNM 86603). Lea, 1852, Obs. Unio, regularly 5: 49. more broadly rounded, somewhat pointed 368 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

or subtruncated. Ventral margin slightly ern creek chub, Semotilus atromaculatus curved, occasionally incurved. Dorsal mar- (Mitchill) (Baker, 1928: 201); both extend drainage. Outline subelliptical, long rhom- over the entire range of S. undulaius (Say). gin straight, ending in a slight angle with Breeding, season. From July to April or the obliquely descending posterior margin. May of the following year. Hinge ligament not very prominent. Pos- Habitat. Lives in small rivers and creeks terior ridge very rounded, but occasionally in mud, sand or gravel. slightly angular with a trace of a secondary Remarks. Strophitus undulatus (Say), ridge above, ending in a slight biangulation is the only species of Strophitus in the near the base. Umbos rather full, located Interior Basin and Atlantic Slope region. in the anterior third of the shell, their The shell has no striking characters and sculpture consisting of a few very strong might be confused with species of Ano- ridges which turn up posteriorly where donta, since members of both genera lack there are a few radial lirae. Surface of the lateral teeth. Anodonta have no pseudo- shell with fine irregular growth lines, gen- cardinals, and do have a straight hinge line; erally smooth and shiny, though sometimes whereas Strophitus have vestigial pseudo- covered with brownish periostracum. When cardinals which appear as slight swellings not heavily covered with periostracum, the anterior to the umbos, and a hinge line shell may be yellowish, greenish or tawny, which is incurved in front of the umbos. often with greenish rays over the entire S. undulatus is replaced in the Alabama- surface or on the posterior slope. Coosa River system and Apalachicolan Each valve has vestiges of pseudo- region by S. subvexus (Conrad) (Johnson, cardinal teeth which are merely slight 1967: 2) which differs from undulatus by swellings, anterior to the umbos. The hinge being distinctly rhomboidal, having a plate is narrow, and slightly incurved in sharper posterior ridge, and by having at front of the umbos. Lateral teeth are ab- least one pseudocardinal in each valve sent. Beak cavities shallow, with muscle elevated slightly above the hinge line. undulatus be confused scars on the hinge plate; anterior adductor Strophitus might with Anodontoides radiatus muscle scars distinct, posterior ones faint. (Conrad) 1967: in these areas, Pallial line scarcely visible. Nacre bluish, (Johnson, 6) except that radiatus is more has broad straw-colored or pinkish toward the umbos. elliptical, green rays, and each valve has a long, Li-imth Height Width mm mm mm narrow pseudocardinal tooth which is elevated to 88 55 24 Swift Creek, 3 mi. SSW slightly and roughly parallel Gainer, Wake Co., North the hinge line. Carolina. Clarke and Berg (1959: 43) have al- 74 44 27 8 Turkey Creek, mi. NW ready discussed the necessity of discarding Edgefield, Edgefield Co., the name Swainson for this species. South Carolina. rugosus Ortmann stated that undulatus 47 29 17 As above. (1919: 195) was only a poorly defined variety of Lea Anatomy. Figured by ( 1838, pi. 15, cdentulus (= rugosus), "connected by fig. 47). Discussed by Ortmann (1911: intergrades," and that they were the same 299). The glochidia are subtriangular, species. He retained the two names, "thus with hooks, and measure 0.36 mm in length avoiding inconvenience." Since only one and 0.30 mm in height. They may either species of Strophitus occurs in the Interior complete their development in the mar- Basin and Atlantic Slope region, it is clear supia of the parent or metamorphose while that, by priority, undulatus must be used. parasitizing the largemouth bass, Micro- Range. Interior Basin: Mississippi and pterus salmoides (Lacepede), or north- Ohio drainages, from Central Texas to Atlantic Slope Unionidae • Johnson 369

Lake Winnipeg, Canada. Atlantic Slope: mann, 1919: 204); all Rockbridge Co. tributary of the upper Savannah River James River, opposite Maidens, Goochland system, South Carolina, north to the St. Co. James River, Cartersville, Cumberland Lawrence River system, Canada. Co.

Specimens Examined York River System South Anna River Savannah River System Drainage. Virginia: [North Fork, S Anna River] Orange, Savannah River Drainage. South Caro- Orange Co. lina: Turkey Creek, 8 mi. NW Edgefield, Edgefield Co. Rappahannock River System Rapiclan River Drainage. Virginia: Cooper-Santee River System Rapidan River, Rapidan, Culpeper Co. Congaree River Drainage. Soutli Caro- (Ortmann, 1919: 204). lina: Congaree River. Rappahannock River Drainage. Vir- ginia: Rappahannock River, Remington, Cape Fear River System Fauquier Co. Mountain Run, Culpeper, Cape Fear River Drainage. North Caro- Culpeper Co. both (Ortmann, 1919: 204). lina: Rocky River, 11 mi. N Sanford, Chat- ham Co. Potomac River System Potomac River Drainage. West Vir- Neuse River System ginia: South Branch Potomac River, Rom- Neuse River Drainage. North Carolina: ney, Hampshire Co. (Ortmann, 1919: 204). Beaverdam Creek, 4.75 mi. S Creedmoor Pennsylvania: Conochocheague Creek, East (57); Crabtree Creek, 6 mi. NW Raleigh Greencastle and Scotland; Branch, both (38); Neuse River, Pooles Bridge; Swift Little Antietam Creek, Waynesboro; 1919: both Franklin Co. Creek, 3 mi. SSW Garner (20); all Wake (Ortmann, 202); Co. Virginia: North Fork, Shenandoah River, E Woodstock, Shenandoah Co. South River, Pamlico River System Waynesboro, Augusta Co. (Ortmann, 1919: South Shenandoah Elk- Tar River Drainage. North Carolina: 204). Fork, River, Co. 1919: Tar River, 2 mi. W Springhope, Nash Co. ton, Rockingham (Ortmann, 204). Maryland: Potomac River, Hancock, Roanoke River System Washington Co. (Ortmann, 1919: 204). Potomac Paton Id., .75 mi. W Point Roanoke River Drainage. Virginia: River, of Frederick Co. District Co- Mason Creek, Salem; Roanoke River, Rocks, of lumbia: Lake. Salem; both (Ortmann, 1919: 204); both Aqueduct Roanoke Co. Subfamily Lampsilinae (Ihering 1901) Chowan River System Ortmann 1910 Genus Carunculina Baker Nottoway River Drainage. Virginia: Continuation of Nottoway River, 3 mi. E Rawlings, Bruns- Toxolasma Rafinesque 1831, Ohio wick Co. Monog. Bivalve Shells River (Phila.), p. 2. Species listed: Unio cyclips; U. cinerescens; U. lividus; U. flexus; all Rafinesque. River System James Type species, Unio lividus Rafinesque. By elimi- Frierson, 1914, Nautilus, 28: 7. James River Drainage. Virginia: Calf- nation, Ortmann and Walker, 1922, Occ. Papers, Mus. River, Goshen 1919: pasture (Ortmann, Zool. Univ. Mich. no. 112, pp. 54, 55, show = 2 mi. 204); North [ Maury] River, WNW that U. lividus is a women duhium and that Lexington; North River, Buena Vista (Ort- therefore Toxolasma must be disregarded. 370 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

1 Corunculina [sic] Simpson 1898, in F. C. Baker, below the medial line, somewhat pointed Bull. Acad. 3(1): 109. Chicago Sci., in males. Ventral margin straight in males. Type species, Unto parvus Barnes. Monotypic. In females marsujoial swelling causes the Corunculina, corrected in die index and on errata to be somewhat convex a little jdos- sheet, and reconfirmed by Simpson, 1900, margin Proc. United States Natl. Mus., 22: 563. terior of the center. Dorsal margin slightly Ortmann, 1912, Ann. Carnegie Mus., 8: 377, curved, forming a distinct angle with partim. (Under Eunjnia.) the obliquely descending posterior margin. Call (1896) monographed Carunculina, Posterior ridge double, sometimes broadly and indicated that it included only a few rounded, but more often both ridges very very variable species, and that most of its angular. Posterior slojoe slightly concave. radiation was in the Interior Basin. He Umbos jorominent, not much elevated probably correctly reduced to synonymy above the hinge line, located in the many of the taxa subsequently recognized anterior third of the shell, their sculjDture by Simpson (1914, 1: 148-161) and Frier- consisting of several concentric sharp son (1927: 87-89). ridges parallel to the growth lines, which curve upward to the posterior ridge. Perios- Carunculina pulla (Conrad) tracum generally with distinct growth Plate 17: 4-7 lines, often satiny, generally blackish, U)iio pullus Conrad 1838, Monography Unionidae, though sometimes brownish, greenish, 2 no. 11, p. 100, pi. 55, fig. (Wateree River, olivish, and with obscure very fine green South Carolina; figured type ANSP [lost]; rays. Warm [= Hot Madison Co.], Springs, Springs, Left valve with two raised North Carolina [Tennessee River system]; prob- triangular in front of the ably either Villosa vancuxemensis (Lea) or rjseudocardinal teeth, one Carunculina glans (Lea) (Ortmann and Walker, other, occasionally crenulate. Hinge line 1922: 55). Conrad (1838: 101) states that short and very narrow before two short, this specimen was seen subsequent to the prepa- straight, lateral teeth. Right valve with ration of the plate. Therefore, the type locality one rather chunky, triangular pseudo- is here restricted to the former locality. ) lateral cavities Lampsilis pulla (Conrad). Simpson, 1914, Cat. cardinal; one tooth. Beak Naiades, 1: 160. shallow, with a few dorsal muscle scars Carunculina Bates Occ. patrickae 1966, Papers, under the hinge plate. Anterior adductor Mus. Zool. Univ. Michigan, no. 646, pp. 1-9, muscle scars well imjDressed, posterior ones 3 text figs., 1 pi. (Savannah River at approxi- faint. Pallial line distinct Nacre mately mile point 134.5 U.S. Army Corps of anteriorly. bluish to iridescent. Engineers Map [ = Johnsons Landing, 10 mi. white, pink purplish, W Allendale, Allendale Co.], on the South Length Height Width Carolina bank; holotype MZUM 85274). mm mm mm Carunculina pulla (Conrad). Johnson, 1967, 32.0 Nautilus, 80: 127, pi. 10, figs. 1-4.

Description. Shell small, seldom reach- ing over 35 mm in length. Outline of female long obovatc, of male, elliptical. Valves subinflated, generally thin, somewhat thickened anteriorly. Anterior end regu- larly rounded; posterior end of females more broadly rounded and subtruncated

1 H. B. Baker (1964, Nautilus, 78: 33) has pointed out that since Simpson contributed noth- ing in the original publication of this genus (under article 51 (c), 1964 edition of the In- ternational Code Zool. Nomen.), the authority for Carunculina must be F. C. Baker. Atlantic Slope Unionidae • Johnson 371

Carunculina pulla (Conrad) cannot be It is likely that the ancestors of C. pulla confused with any other species. The entered the Southern Atlantic Slope region species of Villosa exhibit similar sexual through a commingling of the headwaters dimorphism, but otherwise they are quite of the Apalachicola and Savannah river different from pulla. Villosa delumbis systems. (Conrad) and vibex (Conrad) attain much Range. Southern Atlantic Slope: Alta- larger size, have thinner, generally yellow- maha River system, Georgia, north to the ish, distinctly rayed shells, whereas pulla Neuse River system, North Carolina. is heavier for its size, blackish, and if rayed, only obscurely so. Villosa villosa Specimens Examined Lea on the other is ( ) , hand, broadly rayed, Altamaha RrvER System though the rays are sometimes visible only River in transmitted light. Villosa constricta Ocmulgee Drainage. Georgia: below Lumber Tel- (Conrad) attains much larger size, has a Ocmulgee River, City, fair Co. D. similarly heavy shell, but it is shiny, has (H. Athearn). Altamaha River no distinct posterior ridge, and the females Drainage. Georgia: Mcintosh Co. have a distinct "constriction" in the post- [Altamaha River] Darien, basal region. (USNM). Carunculina most resembles pulla closely Savannah River System Carunculina parva (Barnes) ( —C. paula (Lea) Clench and Turner, 1956: 193) Savannah River Drainage. South Caro- which is found throughout the Interior lina: Savannah River, approximately mile 134.5 of Basin and Apalachicolan region. The um- point (U.S. Army Corps Engineers = 10 mi. Allen- bonal sculpture is similar, but pulla differs Map ) Johnsons Landing, W from parva by having heavy growth lines dale, Allendale Co. and a generally rougher periostracum; but Santee River System more especially it differs by having a sharp posterior ridge, with a second, less promi- Wateree River Drainage. North Caro- nent ridge above it. The posterior ridge is lina: [Headwaters of] Catawba River; usually not present in typical parva, and Pfeiffers Pond; Stewarts Pond; Bissels when it is, it is generally not as acute as Pond; Beaver Creek; all Charlotte, Meck- in pulla. Occasional specimens of C. pulla lenburg Co. (all ANSP). South Carolina-. have an indistinct posterior ridge, as does Wateree River (Conrad). one of the specimens figured. Nevertheless, Cape Fear River System the sharp posterior ridge is so generally present that pulla, with its relatively iso- New Hope River Drainage. North Caro- lated distribution from parva, appears to lina: University Lake, an impoundment on be quite distinct from it. I have pointed Morgan Creek, 1 mi. W Chapel Hill, out elsewhere (Johnson, 1967c) why it was Orange Co. not necessary to create a new taxon for this Neuse River System species. Carunculina parva (Lea) is abundant in Neuse River Drainage. North Carolina: Black Creek, northern Florida, but no Neuse River, Raleigh, Wake Co. (Lea). Carunculina have been reported from either Genus Villosa Frierson the St. Marys or Satilla river systems, which are between Black Creek and the Altamaha Micromya Agassiz 1852, Archiv fur Naturge- 18 47. listed: Unio River system. To the north, no Caruncu- schichte, (1): Species lapillus Say, Margar it ana fahula Lea, M. cur- lina have been found on the Atlantic Slope reyana Lea, non Micromya Rondani 1840 beyond the Neuse River system. (Insecta). 372 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Type species, Unio lapillus Say. Subsequent double in the male; obscured by a slight Indicis Gene- designation, Herrmannsen, 1852, marsupial swelling in the female. Posterior rum Malacozoorum, Supp. et Corr., p. 83. slope slightly concave. Umbos moderately Ortmann, 1912, Ann. Carnegie Mus., 8: 337. swollen, elevated above the partim. slightly hinge located in the anterior of the Villosa Frierson, 1927, Check List North Ameri- line, quarter can Naiades, pp. 11, 80. shell, their sculpture consisting of several Type species, Unio villosus Wright. Original fine, low, slightly double-looped ridges. designation. Surface of the shell with irregular growth lines, smooth and but At this writing, it is impossible to tell occasionally shiny, covered with either or dis- how many species there are in Villosa. usually rough on Frierson (1927: 70-79) includes under tinctly satiny periostracum, especially the Periostracum some- Lampsilis, subgenus Ligumia, many taxa posterior slope. times dark which have been included by other authors subshiny, greenish yellow, under for greenish, or more often brownish black, the Micromya ( —Villosa). Except entire surface of the shell with broad the type species, the taxa listed by Frier- green son '(1927: 80, 81) under Villosa are rays interspersed with narrow ones, some- species of Carunculina. In spite of the state times only visible in transmitted light. of the synonymy of this genus, it is clear Left valve with two delicate pseudo- that the majority of its species occur in the cardinal teeth, one in front of the other, Interior Basin. the anterior one somewhat triangular, the hinder one inclined to be vestigial. Hinge Villosa villosa (Wright)* line short and narrow before two short, Plate 17: 8, 9 straight lateral teeth. Right valve with Unio villosus B. H. 12: Wright 1898, Nautilus, two triangular, narrow, parallel pseudo- 32 (Suwannee River [Luraville], Suwannee Co., cardinals separated by a narrow pit, the Florida; syntype USNM 150503, figured by more anterior tooth some- Simpson, 1900, Proc. Acad. Nat. Sci. Phila., quite vestigial, p. 77, pi. 1, fig. 1, selected as lectotype by times absent; one low lateral tooth. Beak Occ. on 3: Johnson, 1967, Papers Moll., 9, cavities shallow, a few dorsal muscle scars pi. 8, fig. 1). under the hinge plate. Anterior adductor Lampsilis villosus (Wright). Simpson, 1914, Cat. ones Naiades, 1: 143. muscle scars well impressed, posterior Villosa villosa (Wright). Clench and Turner, faint, if visible. Pallial line distinct an- 1956, Bull. Florida State 1: 213, pi. Mus., teriorly. Nacre bluish white, occasionally 4, fig. 2. yellowish white, iridescent, especially pos- Shell in Description. usually small size, teriorly. seldom exceeding 60 mm in length. Outline Length Height Width long, elliptical. Valves subinflated, gener- mm mm mm ally thin and translucent. Anterior end 57 28 18 Suwannee River, Luraville, Suwannee Florida. regularly rounded; posterior end of females Co., Lectotype. Female. slightly more broadly rounded, that of 56 29 20 St. Marys River, Nassau males quite pointed. Ventral margin al- Co., Florida. Female. most always broadly curved except in 51 27 20 Spring Creek, Decatur Co., females where a slight marsupial swelling, Georgia. Male. somewhat posterior of the center, renders Habitat. "Limited to streams it straight or slightly convex. Dorsal mar- spring-fed and clear Clench and Turner gin straight, with a very slight, if noticeable, rivers," angle where it meets the obliquely de- (1956: 214); but their report did not in- scending posterior margin. Hinge ligament clude the rather acid and muddy St. Marys small. Posterior ridge broadly rounded, River. Atlantic Slope Unionidae • Johnson 373

Remarks. Villosa villosa (Wright) bears River, Georgia, J. II. Couper [loaned]). Con- Acad. Sci. ser. a resemblance to Villosa oibex (Conrad), rad, 1850, Jour. Nat. Phila., 2, 1: 275, pi. 37, fig. 2; figured holotype MCZ which is more widely distributed and ex- 178778, purchased from J. II. Couper. tends over the whole of V. villosa, range Unio prevostianus Lea 1852, Trans. Amer. Philos. as both similar sexual di- they show Soc, 10: 269, pi. 19, fig. 24 (Eutowah [Etowah] morphism. In the female of villosa there River [North West], Georgia; figured holotype, C. M. Wheatley collection in ANSP [lost]). is, in general, less tendency for the post- Lea, 1852, Obs. Unio, 5: 25. basal swelling to extend below the ventral Unio nigrinus Lea 1852, Trans. Amer. Philos. Soc, margin, and if inclined to be somewhat 10: 284, pi. 24, fig. 44 (West Florida; figured the is posteriorly pointed, point higher. holotype USNM 86132). Lea, 1852, Obs. Unio, The male is pointed posteriorly as in vibex, 5: 40. Unio but the shell is proportionately longer. gracilior Lea 1856, Proc. Acad. Nat. Sci. 8: 262 Creek V. villosa often has a distinctive roughened Phila., (Buckhead [Burke Co.]; Tobesaufke [Tobesofkee] Creek near Macon periostracum which produces a satiny [Bibb Co.]; both Georgia). Lea, 1858, Jour. luster. Acad. Nat. Sci. Phila., ser. 2, 4: 56, pi. 8, fig. Range. Apalachicolan region: Apalach- 38; figured holotype USNM 85088 [localities not Obs. 6: icola River system, east to the St. Marys separated]. Lea, 1858, Unio, 56. Unio rutilans Lea Proc. Acad. Nat. Sci. River system, Georgia. Peninsular Florida. 1856, Phila., 8: 262 Othcalooga [Oothkalooga] Creek, Gordon Co.; Columbus [Muscogee Co.]; both Specimens Examined Georgia). Lea, 1858, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 59, pi. 9, fig. 41; figured St. Marys River System holotype USNM 85093 from [Oothkalooga] Creek. Lea, 1858, Obs. Unio, 6: 59. St. Marys River Drainage. Georgia: St. Unio subcUipsis Lea 1856, Proc. Acad. Nat. Sci. River, Traders Hill, Charleton Co. Marys Phila., 8: 262 (creeks near Columbus [Mus- St. Nas- (MZUM). Florida: Marys River, cogee Co.], Georgia). Lea, 1858, Jour. Acad. sau Co. (USNM). Nat. Sci. Phila., ser. 2, 4: 62, pi. 10, fig. 44; figured holotype USNM 85095. Lea, 1858, Villosa vibex (Conrad) Obs. Unio, 6: 62. Unio sudus Lea Proc. Acad. Nat. Sci. Plate 17: 10,11 1857, Phila., 9: 170 (Dry Creek, near Columbus [Muscogee Plate 18: 1-3 Co.]; Macon [Bibb Co.]; both Georgia). Lea, Unio vibex Conrad, [May] 1834, New Fresh Water 1859, Jour. Acad. Nat. Sci. Phila., ser. 2, 4: Shells United States, p. 31, pi. 4, fig. 3 (Black 194, pi. 21, fig. 77; figured holotype USNM Warrior River, South of Blount's Spring [Blount 85155 from Dry Creek. Lea, 1859, Obs. Unio,

Co.], Alabama; figured holotype ANSP 56488a ) . 7: 12. Published in May, teste Conrad, 1853, Proc. Unio obfuscus Lea 1857, Proc. Acad. Nat. Sci. Acad. Nat. Sci. Phila., 6: 243. Phila., 9: 172 (Flint River, near Macon [Co.], Unio modioliformis Lea, [August or September] Georgia). Lea, 1859, Jour. Acad. Nat. Sci. 1834, Trans. Amer. Pbilos. Soc, 5: 97, pi. 13, Phila., ser. 2, 4: 197, pi. 22, fig. 80; figured

fig. 40 ( Santee Canal, South Carolina; probable holotype USNM 85089. Lea, 1859, Obs. Unio, figured holotype USNM 85029 [differs slightly 7: 15. from figure]). Lea, 1834, Obs. Unio., 1: 209. Unio dispar Lea 1860, Proc. Acad. Nat. Sci. Published in August or September, teste Lea, Phila., 12: 305 (Columbus [Muscogee Co.], 1854, Proc. Acad. Nat. Sci. Phila., 7: 244. Georgia). Lea, 1860, Jour. Acad. Nat. Sci. Unio exiguns Lea 1840, Proc. Amer. Philos. Soc, Phila., ser. 2, 4: 327, pi. 51, fig. 153; figured

1: 287 ( Cbattaboochee River, near Columbus holotype USNM 85101. Lea, 1860, Obs. Unio, [Muscogee Co.], Georgia). Lea, 1842, Trans. 8: 9. Amer. Philos. Soc, 8: 191, pi. 7, fig. 1; figured Unio averillii Wright 1888, Proc. Acad. Nat. Sci.

holotype USNM 84974. Lea, 1842, Obs. Unio, Phila., 40: 115, pi. 3, fig. 4 (Lake Ashby, Volusia 3: 29. Co., Florida; syntype figured by Simpson, 1892, Unio stagnalis Conrad 1849, Proc. Acad. Nat. Sci. Proc. United States Natl. Mus., 15: 414, pi. 56, Phila., 4: 152 (inhabits mill ponds, Ogeechee fig. 6 [not located]). Lectotype USNM 91142, 374 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Width selected by Johnson, 1967, Occ. Papers on Moll., Lengthmm Heightmm mm 3: 5, pi. 7, fig. 4. 100 48 32 Ochee Creek, about 5 mi. Villosa vibex (Conrad). Clench and Turner, SE Toomsboro, Wilkinson 1956, Bull. Florida State Mus., 1: 209, pi. 4, Co., Georgia. Male. fig- 4. 52 29 17 Mill Race, 2 mi. N Sardis, Description. Shell usually small in size, Burke Co., Georgia. not exceeding 60 mm in length, though Female. 27 As Male. occasionally reaching 100 mm. Outline 48 17 above. Valves subelliptical. subinflated, generally Habitat. Lives in mud or soft sand, thin and translucent. Anterior end regu- particularly where rich in vegetable de- rounded; end of females larly posterior tritus, in small rivers and creeks. more rounded, somewhat broadly pointed Remarks. In the Southern Atlantic Slope in males. Ventral margin straight or region, Villosa vibex ( Conrad ) can be con- curved in often slightly males, slightly fused with Villosa delumbis (Conrad) with arcuate in females. Dorsal straight margin which it is often found living. V. vibex has with a slight, if noticeable, angle very broader, less distinct, green rays which are where it meets the obliquely descending less inclined to be broken by sharp growth small. posterior margin. Hinge ligament rests than those of delumbis. Sexual di- Posterior ridge rounded. Posterior broadly morphism is not as strongly developed in concave, occasionally with slope slightly vibex. The males of both species tend to faint wrinkles and ridges. Umbos moder- be somewhat pointed posteriorly. Females swollen, elevated above the ately slightly of vibex tend to be broadly rounded, hinge line, located in the anterior quarter rendering the shell slightly arcuate, while of the shell, their of sculpture consisting the females of delumbis are greatly in- several fine, low, slightly double-looped flated, with the posterior margin subangu- Surface of the shell rather ridges. generally late dorsally and truncate below. In the but smooth, roughened by periostracum Apalachieolan region, vibex can be con- posteriorly. Periostracum usually subshiny, fused with Villosa lienosa (Conrad) which to greenish yellow, yellowish brown almost is allopatric with V . delumbis. See Remarks black, the entire surface with numerous under Villosa delumbis (Conrad) on p. 376. broad greenish rays, which in darker speci- Range. West Gulf Coastal region, mens can be seen in transmitted light. Alabama-Coosa River system, and Apalach- ieolan Pearl River Mis- Left valve with two delicate pseudo- region: system, east to the River cardinal teeth, one in front of the other, sissippi, Suwanee system, Florida. Peninsular Florida. Southern At- the anterior one somewhat triangular, the lantic Slope: Altamaha River system, Geor- hinder one inclined to be vestigial. Hinge gia, north to the coastal ponds of the Cape line rather long and very narrow before Fear River system, North Carolina. two short, straight lateral teeth. Right valve with two triangular, narrow, parallel Specimens Examined pseudoeardinals separated by a narrow pit, Altamaha River System the more anterior tooth vestigial, sometimes one lateral tooth. Beak cavities absent; Ofimilgee River Drainage. Georgia: shallow, a few dorsal muscle scars under Tobesofkee Creek, near Macon, Bibb Co. the Anterior adductor hinge plate. muscle (USNM). Flat Creek, 2 mi. S Perry, scars well impressed, posterior ones faint, Houston Co. Ruck Creek, 5 mi. NW Hawk- if visible. Pallial line distinct anteriorly. insville; Limestone Creek, 4.3 mi. E Nacre bluish white, sometimes pinkish or Ilawkinsville; Tucsawatchee Creek, 5 mi. purple, iridescent posteriorly. SW Hawkinsville; Cedar Creek, Fountains Atlantic Slope Unioniuak • Joluison 375

Mill, 7 mi. SW Hawkinsville; Mosquito Edisto Biver System Creek, 8 mi. SE Hawkinsville; all Pulaski Edisto River Drainage. SoutJi Caro- Co. 8 mi. Brushy Creek, NW Abbeville, lina: Edisto Biver. Wilcox Co. House Creek, Bowens Mill, 9 mi. N Fitzgerald; Dieksons Creek, 10 mi. Cooper-Santee Biver System NE Fitzgerald; both Ben Hill Co. Wateree River Drainage. South Caro- Little River Drainage. Geor- Ocmulgee lina: Wateree Biver, 2.5 mi. W Camden, Little 7 mi. NE gia: Ocmulgee Biver, Kershaw Co. Cochran, Bleckley Co. Alligator Creek, Santee River Drainage. South Caro- 2 mi. W Alamo, Wheeler Co. Gum Swamp lina: Lake Moultrie, Cross, Berkeley Co. Creek, 1 mi. N McBae, Telfair Co. (USNM). Santee Canal (Lea). Oconee River Drainage. Georgia: Ochee Creek, about 5 mi. SE Toomsboro; Pedee Biver System Turkey Creek, 4 mi. NE Allentown; both Lynches River Drainage. South Caro- Wilkinson Co. Ford Branch, 4 mi. W lina: Lynches Biver, 2 mi. NE Bishopville, Creek, 8 mi. W Dublin; Bocky Dudley; Lee Co. both Laurens Co. Ohoopee River Drainage. Georgia: Waccamaw Biver System Little Ohoopee Biver, 12 mi. NE Wrights- Waccamaw River Drainage. North ville; Little Ohoopee Biver, 2 mi. N Kite; Carolina: Lake Waccamaw, [town of] Biver, 4 mi. S Wrights ville; all Ohoopee Lake Waccamaw, Columbus Co. Johnson Co. Ohoopee Biver, Norristown, Emanuel Co. Cape Fear Biver System Ogeechee Biver System Cape Fear River Drainage. North Caro-

lina: ; Sprunts Pond [not located] ( USNM ) Ogeechee River Drainage. Georgia: stream below Greenfield Mill Pond, Wil- Bocky Comfort Creek, 1 mi. N Louisville; (ANSP); both New Hanover Co. Williamson Swamp Creek, Bartow; Nails mington Creek, 2 mi. S Bartow; Bocky Creek, 2 Villosa delumbis (Conrad) mi. S all Co. Barkcamp Wadley; Jefferson Plate 18:4-8 Creek, 7 mi. E Midville; Mill Creek, 5 mi. Unio delumbis Conrad New Fresh Water E Midville; Buckhead Creek, 4 mi. E 1834, Shells United States, p. 35, pi. 5, fig. 3 (small Vidette; Creek, 5 mi. SW Waynes- Bocky streams near Cooper River, South Carolina; all Burke Co. Mill 8 boro; Chew Creek, figured type not in ANSP [lost]). mi. W Millen, Jenkins Co. Unio tenerus Ravenel 1834, Cat. Recent Shells Cabinet Edmund Ravenel, p. 7 (Cooper River, Savannah Biver System South Carolina) [nomen nudum]. Simpson 1892, Proe. United States Natl. Mus., 15: 416, Savannah River Georgia: Drainage. pi. 58, figs. 5, 8 (South Carolina; figured syn- Brier Creek, 7.5 mi. NE Wrens, Jefferson type USNM 85030 [male]; [Savannah River], Co. Mill Bace, 2 mi. N Sardis; Beaverdam Georgia; figured syntype USNM 85032 Mus. Creek, 7 mi. NW Girard; both Burke Co. [female]). Walker, 1919, Oec. Papers, Zool. Univ. Mich. no. 74, p. 1, pi. 1, figs. 1-6. COMBAHEE BlVER SYSTEM Unio vaughanianus Lea 1838, Trans. Amer. Philos. Soc, 6: 5, pi. 3, fig. 5 (Sawney's Creek, near Salkehatchee River Drainage. South [about 8 mi. NW] Camden [Kershaw Co.], Carolina: Little Salkehatchee Biver, 4 mi. South Carolina; two syntypes USNM 86106, smaller than the the other N Ehrhardt, Bamberg Co. Salkehatchee one a bit figure, with the outline of the figure, but with less Biver, Broxton Bridge; Whippy Swamp prominent rays. The figure appears to be a 2.5 mi. both Creek, NE Crocketville; composite.) Lea, 1838, Obs. Unio, 2: 5. Hampton Co. Unio ogeecheensis Conrad 1849, Proc. Acad. Nat. 376 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Sci. Phila., 4: 153 (Ogeechee River, Georgia, traeum often subshiny, greenish yellow, T. H. Conrad, 1850, Couper [loaned]). Jour. brownish, or greenish, the entire surface Acad. Nat. Sei. Phila., ser. 2, 1: 275, pi. 37, with narrow, sometimes very narrow, figs. 3-4; figured syntype, fig. 3, MCZ 146971 which are broken purchased from J. H. Couper, selected as lecto- greenish rays by growth type by Johnson, 1956, Bull. Mus. Comp. Zool., rests to form concentric bands. 115: 126. Left valve with two delicate pseudo- Unio concavus Lea Trans. Amer. Philos. 1S52, cardinal teeth, one in front of the other, the Soc, 10: 260, pi. 15, fig. 11 (Abbeville District anterior one somewhat triangular, the [Savannah River drainage], South Carolina; hinder one inclined to be figured holotype USNM 85154). Lea, 1852, vestigial. Hinge Obs. Unio, 5: 16. line rather long and very narrow before Unio proximus Lea 1852, Proc. Amer. Philos. Soc, two short, straight lateral teeth. Right valve 5: 252 (Georgia). Lea, 1852, Trans. Amer. with two triangular, narrow, parallel pseudo- Philos. Soc, 10: 271, pi. 20, fig. 27; figured cardinals separated by a narrow pit; the holotype USNM 85131. Lea, 1852, Obs. Unio, anterior 5: 27. more tooth vestigial, sometimes cavities Unio contiguus Lea 1861, Proc. Acad. Nat. Sci. absent; one lateral tooth. Beak Phila., 13: 392 (Stewarts Mill Dam, Union Co., shallow, a few dorsal muscle scars under North Acad. Nat. Carolina). Lea, 1862, Jour. the hinge plate. Anterior adductor muscle Sci. Phila., ser. 2, 5: 199, pi. 28, fig. 268; scars well impressed, posterior ones faint, figured holotype USNM 85120. Lea, 1863, if visible. Pallial line distinct Obs. Unio, 9: 21. anteriorly. Nacre sometimes white or Unio genuinus Lea 1868, Proc. Acad. Nat. Sci. dirty pinkish, Phila., 20: 161 (Bissels Pond, Charlotte [Meck- but usually bluish white and iridescent. lenburg Co.] North Carolina). Lea, 1868, Jour. Length Height Width Acad. Nat. Sci. Phila., ser. 2, 6: 305, pi. 46; mm mm mm fig. 117, figured holotype USNM 85123. Lea, 64 39 30 Ogeechee River, Georgia. 1869, Obs. Unio, 12: 64. Lectotype of U. ogeeche- ensis Conrad. Female. Description. Shell usually small in size, 57 35 19 Mill Race, 2 mi. N Sardis, seldom exceeding 60 mm in length. Outline Burke Co., Georgia. of female of male long obovate; long ellip- Female. tical. Valves thin subinflated, generally 53 30 19 As above. Male. and translucent. Anterior end regularly Habitat. Lives in mud or soft rounded; posterior end of females more sand, where rich in broadly rounded and subtruncated below particularly vegetable in small rivers and creeks. the medial line, somewhat pointed in males. detritus, Remarks. In the Southern Atlantic Ventral margin straight in males, in females Slope Villosa delumbis can marsupial swelling causes the margin to be region, (Conrad) be confused with Villosa vibex somewhat convex a little posterior of the only (Con- under which see: Remarks on 374. center. Dorsal margin straight forming a rad) p. Villosa delumbis is close sharp angle with the obliquely descending (Conrad) very to Villosa lienosa Conrad of the posterior margin. Posterior ridge broadly ( ) Apalach- icolan rounded. Posterior slope slightly concave, region. Both species are similarly occasionally with faint ridges and wrinkles. shaped and the females show the same Umbos moderately swollen, slightly ele- sexual dimorphism. V. lienosa has a much vated above the hinge line, located in the heavier shell; the posterior ridge, while anterior third of the shell, their sculpture rounded, tends to be faintly biangulate. consisting of several fine, low, slightly The periostraeum is usually black or very double-looped ridges. Surface of the shell dark yellowish green, and when visible the generally rather smooth and shiny, but rays are solid green. The nacre is often roughened by periostraeum posteriorly pinkish or coppery. V. delumbis has a especially on the posterior slope. Perios- thinner shell, there is scarcely ever a hint Atlantic Slope Uniontoae • Johnson 377

of the posterior ridge 1 being faintly bi- both Burke Co. Chew Mill Creek, 8 mi. angulate, the periostracum is usually a light W Millen, Jenkins Co. Ogeechee River, yellow or light greenish yellow, and the Scarboro; Ogeechee River, bridge 1 mi. rays are almost always distinct and char- 5 Dover; Ogeechee River, 1.5 mi. SW acteristically broken by growth rests. The Oliver; all Screven Co. (all MZUM). nacre is very seldom pinkish. Ogeechee River, bridge 1.5 mi. E Blitch- Although the figured type of V. delumbis ton; Ogeechee River, Jinks Bridge [Rte. has been lost, the life size figure, with its 16]; Ogeechee River, Morgan Bridge, 14 distinct, narrow rays, clearly distinguishes mi. SE Pembroke; Ogeechee River, mouth it from vibex, and it corresponds almost of Arnold Lake; Jones Lake; all Brvan Co. exactly in outline with the male shell (all MZUM). figured in this report. Savannah River System Range. Southern Atlantic Slope: Alta- maha River system, Georgia, north to the Broad River Drainage. Georgia: Broad Neuse River system, North Carolina. River, Elbert, Huguenot Co. (USNM) Broad River, Lincoln Co. Specimens Examined Savannah River Drainage. Georgia: Altamaha River System Savannah River, Augusta, Richmond Co. South Carolina: Savannah River, 2 mi. River Georgia: Ocmulgee Drainage. SW Millettville (MZUM); Savannah River, Macon, Bibb Co. Echeconnee Creek, Eche- Johnsons Landing, 10 mi. W Allendale; connee, Houston Co. Buck Creek, 5 mi. both Allendale Co. Georgia: Savannah NW Hawkinsville; Limestone Creek, 4.3 River, 7.5 mi. NE Shell Bluff; Brier Creek, mi. E Hawkinsville; Tuesawhatchee Creek, 6 mi. N Waynesboro; outlet of Waynesboro 5 mi. SW Hawkinsville; Cedar Creek, 8 mi. Lake, Waynesboro; Mill Race, 2 mi. N Sar- SW Hawkinsville; all Pulaski Co. dis; all Burke Co. Oconee River Drainage. Georgia: Ochee Creek, about 5 mi. SE Toomsboro; Combahee River System Turkey Creek, 4 mi. NE Allentown; both Salkehatchee River Drainage. South Wilkinson Co. Ford Branch, 4 mi. W Dub- Carolina: Pauline Cave, near Kline, Barn- lin; Palmetto Creek, 7 mi. S Dublin; both well Co. Laurens Co. (MZUM). Ohoopee River Drainage. Georgia: Edisto River System Ohoopee River, 4 mi. S Wrightsville; Little Edisto River South Caro- Ohoopee River, 1 mi. E Kite; both Johnson Drainage. lina: Snake Co. Co. Ohoopee River, 1 mi. E Adrian; Swamp Creek, Orangeburg 6 mi. Co. Edisto Ohoopee River, Norristown; Yam Grande NE Bamberg, Bamberg Creek, 3 mi. W Swainsboro; all Emanuel River, Canadys Landing, 8 mi. SW St. Co. George, Dorchester Co. Edisto River, 1 Altamaha River Drainage. Georgia: mi. E Jacksonboro, Colleton Co. (MZUM). Darien, Mcintosh Co. (USNM). Cooper-Santee River System Ogeechee River System Congaree River Drainage. South Caro- Ogeechee River Drainage. Georgia: lina: Congaree River. Ogeechee River, Shoals, Warren Co. Catawba River Drainage. North Caro- Ogeechee River, 4 mi. SW Mitchell, Glas- lina: Long Creek; Catawba River; Bissels cock Co. Williamson Swamp Creek, Bar- Pond, Charlotte; Pfeiffers Pond, Charlotte tow, Jefferson Co. Barkcamp Creek, 7 mi. (ANSP); Flannigans Pond, Charlotte E Midville; Mill Creek, 5 mi. E Midville; (ANSP); all Mecklenburg Co. [Ten Mile 378 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

that it was from Creek] of Catawba River, Wolfsville [7 mi. specimen was figured, Deep Chatham North and W Monroe], Union Co. (MZUM). River, Gulf, Co., Carolina, the so restricted. Wateree River South Caro- type locality is, here, ) Lea, Drainage. 1862, Obs. Unio, 8: 61. 8 mi. lina: Sawneys Creek, about NW Lampsilis constricta (Conrad). Simpson, 1914, Camden, Kershaw Co. (USNM). Cat. Naiades, 1: 111. Cooper River Drainage. South Caro- Eunjnia (Micromya) constricta (Conrad). Ort- mann, 1915, Nautilus, 29: 66. lina: near Cooper River (Conrad). Shell small in seldom Pedee River System Description. size, exceeding 50 mm in length. Outline of North Caro- Yadkin River Drainage. female short obovate; of male subelliptical. lina: Stewarts Mill Dam, Union Co. Valves not much inflated, rather heavy and (USNM). strong. Anterior end regularly rounded; posterior end of females more broadly Waccamaw River System rounded and pointed a little more than Waccamaw River Drainage. North midway up from the base, truncated below Carolina: drainage canal beside Lake the medial line. In mature specimens there Waccamaw, 1 mi. NNW Dupree Landing, is a rather distinct "constriction" in the Columbus Co. middle of this truncation which marks the boundary between the branchial opening Cape Fear River System and the papillar part of the mantle edge Cape Fear River Drainage. North Caro- in front of it; somewhat pointed in males, lina: University Lake, an impoundment on the point being generally less than midway Morgan Creek, 1 mi. W Chapel Hill, up from the base. Ventral margin quite Orange Co. Rocky River, 11 mi. N Sanford, convex in females, slightly convex in males. Chatham Co. (MZUM). North East Cape Dorsal margin rather rounded, forming a Fear River, Dublin Co. (MZUM). Green- more or less distinct angle with the field Mill Pond, Wilmington, New Hanover obliquely descending posterior margin. Co. (USNM). Posterior ridge broadly rounded. Posterior slope slightly concave, occasionally with River System Neuse fine radiating wrinkles. Umbos not much Neuse River Drainage. North Carolina: swollen, slightly elevated above the hinge of the North Flat River, 5.5 mi. S Roxboro Center, line, located in the anterior quarter their of Person Co. (81). Neuse River (USNM). shell, sculpture consisting evenly double-looped ridges. Surface of the shell Villosa constricta (Conrad) generally rather smooth and shiny, some- Plate 18: 9-10 times roughened with concentric growth lines. Periostracum often subshiny, yellow- Unio lienosus constrictus Conrad 1838, Mono- ish or bottle graphy Unionidae North America, no. 10, p. 91, green, green, becoming brownish with with pi. 49, fig. 4 ( North [ = Maury] River, Rock- age, generally distinct, bridge Co., Virginia; figured holotype ANSP but very fine, green rays over the entire 20423, now ANSP 56465a.) surface. Unio Lea Proc. Acad. Nat. Sci. genthii 1857, Left valve with two rather heavy pseudo- Phila., 9: 85 (Catawba River, Caston Co.; Deep cardinal teeth, one in front of the other, River, Gulf [Chatham Co.]; both North Caro- both rather line lina). Lea, 1862, Jour. Acad. Nat. Sci. Phila., triangular. Hinge very ser. 2, 5: 57, pi. 2, fig. 204; figured holotype short and very narrow before two straight Roanoke USNM 84834, labeled, River, Raleigh, lateral teeth. Right valve with two pseudo- North Carolina, Prof. Emmons. (This locality cardinals, the posterior one chunky and is nonexistent. There is no way to tell which serrated, the more anterior tooth vestigial; locality the type was from; so it is assumed here that since it was probably Prof. Emmons' one lateral tooth. Beak cavities shallow, Atlantic Slope Unionidae • Johnson 379

a few dorsal muscle scars under the hinge posterior ridge and a generally smooth, plate. Anterior adductor muscle scars well shiny surface that is distinctly rayed. impressed, posterior ones faint, if visible. Villosa constricta (Conrad) is allopatric Pallial line generally distinct anteriorly. with Villosa vanuxemensis (Lea) (Simp- Nacre sometimes dirty white or yellowish, son, 1914, 1: 105) of the Cumberland and especially toward the beak cavities, but Tennessee rivers. The constriction found usually bluish white and iridescent. rather regularly in old females of con- stricta is not as well in vanuxe- Length Height Width developed mm mm mm mensis, and the latter has a distinct nacre 49.0 28.0 17.0 Pedlar River, 1 mi. W color (usually an unreliable character) Pleasantview, Amherst which varies from dirty purplish white Co., Virginia. Male. through salmon-tinted to dark purple, usu- 41.4 27.0 16.0 North River, Rockbridge ally darker toward the beak of the Co., Virginia. Female. cavity Holotvpe of Unto con- shell, whereas the nacre color of constricta strictus Conrad. is dirty white, bluish white or yellowish 39.0 27.0 18.0 [Deep River, Gulf, Chat- toward the beak cavities. Ortmann (1915: ham North Caro- Co.], 66) collected a number of specimens of lina. Male. Holotype of vanuxemensis on September 17, 1912 in the Unio genthii Lea. North Fork, Holston River, Saltville, Smyth season. season Breeding The breeding Co., Virginia, and at once recognized its in and ends in Glo- begins August June. affinity with constricta. chidia than subspatulate, higher long. Range. Southern Atlantic Slope: Length 0.21 mm, height 0.27 mm (Ort- Catawba River, North Carolina, of the mann, 1915: 66, 67). upper Cooper-Santee River system, north Habitat. Lives in sand in rather swift, to the James River system, Virginia. flowing rivers. Remarks. Villosa constricta (Conrad) of Specimens Examined the Southern Atlantic Slope region does not Cooper-Santee River System much resemble any other unionid found there. There is often quite a bit of differ- Catawba River Drainage. North Caro- ence in the degree of inflation and form lina: Catawba River, Gaston Co. (Lea). of the shell, but it differs from all of the Beaver Creek [Gaston Co.] (MZUM). other Atlantic Slope Villosa in the shape Long Creek [Mecklenburg Co.] (MZUM). of the postbasal swelling which generally [Ten Mile Creek] of Catawba River, does not project beyond the base line of Wolfsville [7 mi. W Monroe], Union Co. the shell, and in the emargination behind (USNM). the swelling. For its size constricta is much Cape Fear RrvER System heavier and stronger than any other Villosa in the region under study, and has a dis- Deep River Drainage. North Carolina: tinctive yellowish green or shiny dark Deep River, Gulf, Chatham Co. (Lea). chestnut-brown periostraeum, and the fe- Cape Fear River Drainage. North Caro- male has a distinct constriction in the lina: Rocky River, 11 mi. N Sanford, Chat- postbasal region. Females of Caninculina ham Co. (MZUM). pulla (Conrad) show a similar sexual Neuse RrvER System dimorphism, but they lack the constriction. In addition, pulla usually has a sharp Neuse River Drainage. North Carolina: double posterior ridge, a rough periostra- North Flat River, 5.5 mi. S Roxboro, Person eum and if rayed at all, obscurely so, Co. (81). Eno River, 1.8 mi. S Cedar whereas constricta has a Grove 122 both broadly rounded ( ) ; Eno River, Hillsboro; 380 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Orange Co. Eno River, 6 mi. NW Durham Ligumia contains two species, L. recta

(113); Fork, Little River, 11 mi. NNW (Lamarck) (Ortmann, 1919: 276, pi. 16, Durham 101 both Durham Co. Neuse so-called latis- ( ) ; figs. 12, 13) (the subspecies River, 6 mi. E Raleigh; Hare-Snipe Creek, sima (Rafinesque) is an ecophenotype, oc- 6.25 mi. NW Raleigh (37); Little River, curring in rivers) and L. nasuta (Say). Tarpleys Mill, 2 mi. NE Wendell (5); all Wake Co. Ligumia nasuta (Say) Plate 19: 1,2 Pamlico River System Unio nasutus Say 1817, Nicholson's Encyclopedia, 2 Tar River Drainage. North Carolina: [no pagination], pi. 4, fig. 1 (Delaware and Tar River, 2 mi. ENE Bunn, Franklin Co. Schuylkill [rivers near Philadelphia, Philadel- phia Co., Pennsylvania]; ANSP Chicod Creek, Pitt Co. (MZUM). type, [lost]). Ohiiquaria attentuata Rafinesque 1820, Ann. Gen. des Sci. 5: 304 Roanoke River System Physiques (Bruxelles), (la fleuve Hudson [New York]; tvpe, not in ANSP Staunton River Drainage. Virginia: [lost]). Unio rostrata Valenciennes in Humboldt Mason Creek, near Roanoke River, Salem; 1827, and Bonpland, Voyage aux Regions Equinoxi- Tinker Creek, Roanoke both (MZUM); ales du Nouveau Continent, pt. 2, 2: 233, Roanoke Co. pi. 53, fig. 3 (Philadelphia vicinis [near] [Philadelphia Co., Pennsylvania]; type [location James River System unknown] ). Unio vaughanianus Sowerby 1868, Conch. Iconica, James River Drainage. Virginia: Dun- 16, Unio, pi. 61, fig. 308, non Lea, 1838. lap Creek (MZUM); Jackson River, Cov- Unio fisherianus Kuester 1860, Conch. Cabinet, ington (MZUM); Cowpasture River, near 9, pt. 2, pi. 68, fig. 6, non Lea, 1838. Longdale (MZUM); all Alleghany Co. Lampsilis nasuta (Say). Simpson, 1914, Cat. Naiades, 1: 97. Calfpasture River, Goshen (MZUM); = Eurynia nasuta (Say). Ortmann, 1919, Mem. North [ River, 2 mi. WNW Maury] Carnegie Mus., 8: 271, pi. 16, figs. 10, 11. Lexington; both Rockbridge Co. James Ligumia nasuta (Say). Clarke and Berg, 1959, River, Buchanan, Botetourt Co. Pedlar Cornell Univ. Exp. Sta. Mem. 367, p. 51, 54. River, 1 mi. W Pleasantview, Amherst Co. fig. River, Beaver Creek, 6 mi. James opposite Description. Shell generally medium in E Co. Lynchburg, Campbell (MZUM). size throughout most of its range, usually Cumberland Co. James River, Cartersville, not exceeding 80 mm in length, though James River, opposite Maidens, Goochland reaching over 110 mm in a few localities. Co. River, Henrico Co. James Richmond, Outline elongated, subelliptical, or sub- (ANSP). lanceolate; distinctly over twice as long as high. Valves usually subcompressed, thin Genus Ligumia Swainson to subsolid, inequilateral. Anterior end Ligumia Swainson Treatise on 1840, Malaeology, regularly rounded; posterior end elongated pp. 268, 274, 378. and bluntly pointed. Ventral margin Type species, [Unio] recta Lamarck. Original curved. Dorsal designation, p. 274. margin straight, slightly Ortmann, 1912, Ann. Carnegie Mns., 8: 338, curved, forming a distinct angle, at least partim. in immatures, with the obliquely descend- Ortmann and Walker, 1922, Occ. Papers, Mns. ing posterior slope. Hinge ligament rather Zool. Univ. Midi., no. 112, p. 59. long and low. Posterior ridge distinct and Frierson (1927: 70-79) includes many angular, becoming toward the umbos taxa in this genus. Most of them have been broader, more rounded, and indistinct. placed by subsequent authors under Lam))- Posterior slope slightly concave, near the silis s. s. and Villosa. It appears that umbos becoming flatter or even somewhat Atlantic Slope Qniontdae • Johnson 3S1

convex posteriorly. Umbos very low, lo- Lake Erie that this species breeds from cated in the anterior quarter of the shell, August until the following June. their sculpture consisting of fine close Habitat. Lives in sand and mud, mostly ridges which are looped in front and run in protected areas in ponds, lakes, canals, parallel to the axis of the shell behind. and in slack water areas of streams. Surface of the shell with irregular growth Remarks. Ligumia nasuta (Say) of the lines, sometimes with a few nearly vertical upper Ohio River drainage and Atlantic- ridges below the posterior ridge. Perios- Slope region is distinguishable from any tracum usually fine, subshiny, greenish other North American unionid by its yellow, dark olive to brownish, often with usually distinctly lance-head shaped pos- distinct fine rays, especially posteriorly. terior end. On the Atlantic Slope the male Left valve with two compressed pseudo- shell can be confused with Elliptio lanceo- cardinal teeth, one in front of the other, lata (Lea), but the latter does not have both subtriangular and crenulated; no the same distinct pattern of rays, nor the interdentum; two long straight laterals. dull olive green color. Also the nacre of Right valve with two pseudocardinals, the nasnta is generally silvery white or cream posterior one triangular, the more anterior colored, whereas that of lanceolata may one low and vestigial; one lateral tooth. be coppery or purple. The females of Beak cavities very shallow, with a few nasuta have a distinct expansion in the dorsal muscle scars. Anterior adductor postbasal region which render them more muscle scars well impressed; posterior ones easily distinguishable from lanceolata than often scarcely visible. Pallial line distinct the male shell. anteriorly. Nacre bluish white, often cream Clarke and Berg (1959: 51), apparently color or toward the beak 1 : 97 the salmon, especially following Simpson ( 1914, ) , give cavity, posteriorly iridescent. southern range of this species as North Sexual differences well marked in the Carolina. Simpson's record appears to be shell. The male shell tapers uniformly based on series of rather posteriorly pro- behind into a point, but the female is dis- duced specimens of Elliptio lanceolata tinctly produced in the postbasal region, (Lea) from Wilmington, New Hanover behind which the ventral margin slopes Co., North Carolina. Ligumia nasuta is up more acutely than in the male, and is decidedly part of the northern element of straight or concave. the Atlantic Slope fauna, the James River, Width Virginia, being its southernmost record. Lengthmm Heightmm mm (Conrad, 1836: 38, pi. 18, fig. 1, figures a 112 44 28 Connecticut River, Hart- female specimen from the James River, but ford, Hartford Co., Con- necticut. Male. it has not been collected there since his 66 29 14 Potomac River, Washing- time.) As observed by Conrad, nasuta ton, District of Columbia. does not go very far up into the rivers in Female. the southern part of its range, and we are 62 25 12 As above. Male. led by Conrad to believe that his record Anatomy. Discussed by Ortmann (1911: from the James is from the tidewater 343); discussed and figured by Reardon region. It is abundant in the Potomac is not (1929: 7, pi. 2, figs. 1-10). The glochidia River in the tidewater region, but are subovate, with an undulate hinge line, found very far up the river. Ortmann and measure 0.25 mm in length and 0.29 (1919: 275) found the distribution to be mm in height. The host fish is unknown. similar in Pennsylvania. Breeding, season. Ortmann (1919: 272) Range. Atlantic Slope: James River indicates from specimens he examined in system, Virginia, north to the St. Lawrence 382 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

River system, Canada, westward through lampsilis (Lampsilis) cariosa (Say) the Mohawk River and Erie [now New Plate 19:3-5 York State both New York. Barge] Canal, Unio cariosus Say 1817, Nicholson's Encyclopedia, Interior Basin: Lake Erie, Ohio, and 2 [no pagination], pi. 3, fig. 2 (Delaware and Michigan. Schuylkill Rivers; [Susquehanna River], Wilkes Rarre; [Luzerne Co.; all Pennsylvania]; type, Specimens Examined ANSP [lost]). The type locality was restricted to: Schuylkill River, near Philadelphia, Penn- James River System sylvania, and a neotype MCZ 178839 was selected by Johnson ( 1947, Occ. Papers on James River Drainage. Virgino: James Moll, 1: 148, pi. 19, fig. 1). River (Conrad). Lampsilis pallida Rafinesque 1820, Ann. Gen. des Sci. Physiques (Bruxelles), 5: 299 (Hudson Potomac River System River, [New York]; type not in ANSP [lost]). Unio ovata Valenciennes 1827, in Humboldt and Potomac River District Drainage. of Bonpland, Voyage aux Regions Equinoxiales du River. Columbia: Potomac Nouveau Continent, pt. 2, 2: 226, pi. 50, figs. 1 a-c, (pres de Washington [District of Co- non Genus Lampsilis Rafinesque lumbia]; type [location unknown]) Say 1817. Lampsilis Rafinesque 1820, Ann. Gen. des Sci. Unio crocatus Lea 1841, Proc. Amer. Philos. Soc, Physiques (Bruxelles), 5: 298. Species listed: 2: 31 (Savannah River, Georgia). Lea, 1842, Lampsilis cardium Rafinesque, Lampsilis ovata Trans. Amer. Philos. Soc, 8: 238, pi. 22, fig. (Say), Lampsilis fasciola Rafinesque. 52; figured holotype USNM 84908. The lecto- Type species, Unio ovatus Say. Subsequent type selected by Johnson ( 1947, Occ. Papers designation, 1847, Indicis Gene- Herrmannsen, on Moll., 1: 156) but not figured, is invalid. rum 1: 575. Malacozoorum, Ortmann, 1912, The holotype which was catalogued under the Ann. 8: 345. Carnegie Mus., same number has since been located. Lea, Aeglia Swainson 1840, Treatise on Malacology, 1842, Obs. Unio, 3: 76. 378. listed: Ac. ovata pp. 265, Species (Say), Unio oratus Conrad 1849, Proc. Acad. Nat. Sci.

occidens Lea ) . ( Phila., 4: 153 (Flint River, Georgia; here re- Unio ovatus 1 Type species, Say. Subsequent stricted to: Ogeechee River, Georgia ). Conrad Indicis designation, Herrmannsen, 1846, Gene- 1850, Jour. Acad. Nat. Sci. Phila., ser. 2, 1: 267, rum 1: 20. Malacozoorum, pi. 37, fig. 6; type, coll. of J. H. Couper [lost]. Lampsilis cariosa (Say). Simpson, 1914, Cat. Subgenus Lampsilis s.s. Naiades, 1: 43. Ortmann, 1919, Mem. Carnegie Mus., 8: 313, pi. 20, figs. 3-5. Johnson, 1947, All of the species described in this paper Occ. Papers Moll., 1: 145, pi. 19, figs. 1-2. belong to Lampsilis s. s. Frierson (1927: Clarke and Berg, 1959, Cornell Univ. Exp. Sta. 67-86) lists ten other subgenera, three of Mem., no. 367, p. 54, figs. 59-60. Clarke and Natl. Mus. Bull. no. which are of his own creation. One of Rick, 1963, Canada, 189, p. 27, pi. 1, fig. 3. them, Villosa, is now used as a replace- Shell ment name for Micromya Agassiz (see under Description. usually medium, 130 mm in Out- Villosa on p. 371. Ligumia Swainson now though reaching length. line of male and somewhat elon- replaces Eurynia Rafinesque. To comment elliptical of female subovate, or obovate, rather on the other subgenera is not in the scope gate; short and Valves some- of this paper, but on cursory examination, high. inequilateral, the present author disagrees substantially 1 Also described on the same in both with Frierson's classification, both on a page publications is Unio contrarius = Lampsilis clai- generic and specific level. bornensis ( Lea ) with Ogeechee River, Georgia, Lampsilis s. s., while clearly of Interior as the type locality. The localities for the two

1 are since oratus is Basin origin, appears to have speciated species obviously transposed, not found in the Flint River, but in the Ogeechee about equally there and in the combined River, and contrarius is found in the Flint River Apalachicolan and Atlantic Slope regions. and not in the Ogeechee River. Atlantic Slofe Umontdae • Johnson 383

what inflated; moderately heavy. Anterior end regularly rounded; posterior end more broadly rounded and somewhat pointed in the male, truncated in the female. Ven- tral margin slightly curved. Dorsal margin straight, terminating in a broad angle with the obliquely descending posterior margin. Hinge ligament prominent, running under the umbos and appearing anteriorly. Pos- terior ridge rounded and poorly defined. Posterior slope slightly convex, usually with a few faint ridges and wrinkles. Umbos moderately swollen, slightly raised above the hinge line, located somewhat anterior to the middle of the shell, their sculpture consisting of four or five wavy recurved ridges. Surface of the shell generally smooth except for a few growth ridges. Periostracum usually shiny, bright wax or straw 7 yellow , infrequently greenish yellow, becoming a dirty brownish yellow or red- dish brown in matures. Green or blackish rays, when present, usually restricted to the posterior slope. Left valve with two compressed, ser- rated pseudocardinal teeth, one in front of the other, the anterior one somewhat tri- angular, the hinder one lower, but chunky, located directly under the umbo. Hinge line rounded, with a considerable inter- dentum under the umbo before two short lateral teeth. Right valve with two oppos- ing pseudocardinals, the anterior one low and laminate, the other higher, chunky and serrated; one lateral tooth, sharply truncated behind. Beak cavities moder- ately deep, with deep dorsal muscle scars. Anterior adductor muscle scars well im- pressed, posterior ones less so. Pallial line distinct. Nacre bluish white or tinged with salmon.

Width Lengthmm Heightmm mm 30 384 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Range. Atlantic Slope: Ogeechee River Landing, 2 mi. W Murrells Inlet, George- system, Georgia, north to the Sydney River, town Co. Nova Scotia (Clarke and Rick, 1963: 27), Cape Fear River System and in the St. Lawrence River system, Canada, westward to the Madawaska Cape Fear River Drainage. North Caro- River, Ontario, of the lower Ottawa River lina: Cape Fear River, Cumberland Co. drainage. (USNM).

Specimens Examined Neuse RrvER System Neuse River North Carolina: Ogeechee River System Drainage. Eno River, 6 mi. NNW Durham, Durham River Ogeechee Drainage. Georgia: Co. (112). Neuse River [6 mi. E] Raleigh; Warren Co. Ogeechee River, Shoals, tributary of Swift Creek, 5 mi. SW Raleigh Screven Co. Ogeechee River, Scarboro, (22); both Wake Co. (MZUM). Ogeechee River, mouth of Arnold Lake [Rryan Co.] (MZUM). Pamlico River System Tar River North Carolina: Savannah River System Drainage. Tar River, 2 mi. W Springhope, Nash Co. Savannah River Drainage. South Caro- River System lina: Abbeville District (USNM). Edge- Chowan field District. Savannah 2 mi. River, SW Nottoway River Drainage. Virginia: Millettville Savannah (ANSP); River, Nottoway River, 3 mi. E Rawlings, Bruns- Johnsons Landing, 10 mi. W Allendale; wick Co. Savannah River, Kingjaw Point, 10 mi. WSW Allendale (ANSP); all Allendale York River System Co. Savannah River, 7.5 mi. NE Georgia: Pamunkey River Drainage. Virginia: Shell Savannah 6 mi. NE Rluff; River, Pamunkey River, 2 mi. N Hanover, Han- Girard; both Rurke Co. Relow Haga Slage over Co. Landing, E Sylvania (ANSP); Savannah Potomac River System River, 7 mi. NE Newington; both Screven Co. Savannah River, in sandbank, Savan- Potomac River Drainage. Maryland: nah, Chatham Co. (MZUM). Monocacy River, 2 mi. W Mt. Pleasant, Frederick Co. Potomac River, Cabin John, Cooper-Santee Rpver System Montgomery Co. (Ortmann, 1919: 316). Potomac Great Fairfax Co. Congaree River Drainage. South Caro- River, Falls, District Columbia: Potomac River. lina: Congaree River. of Wateree River Drainage. South Caro- Lampsilis (Lampsilis) dolabraeformis (Lea) lina: Wateree River, 2.5 mi. W Camden, Plate 20: 1-4 Kershaw Co. Unto dolabraeformis Lea 1838, Trans. Araer. Santee River Drainage. South Caro- Philos. Soc, 6: 103, pi. 24, fig. 113 (Altamaha lina: Santee River, Wilsons 5 mi. Landing, River, Liberty [now Long] Co.; Altamaha NW Pineville (ANSP); below Santee River River, near Darien [Mcintosh Co]; both Dam, near St. Stephens; both Berkeley Georgia; figured holotype USNM 84888). Lea, Co. 1838, Obs. Unio, 2: 103. Lampsilis dolabraeformis (Lea). Simpson, 1914, Waccamaw River System Cat. Naiades, 1: 46.

Waccamaw River Drainage. South Description. Shell large, often exceed- Carolina: Waccamaw River, Wachasaw ing 130 mm in length. Outline of male Atlantic Slope Unioxidae • Johnson 385

somewhat of fe- Length Height Width elliptical and elongate; mm nun mm male obovate, rather short and Valves high. 129 91 64 Altamaha River, 4 mi. N Anterior inflated, rather heavy. end regu- Jesup, Wayne Co., Georgia. larly rounded; posterior end more broadly Male. rounded and somewhat pointed in the 108 78 55 As above. Female. male, truncated in the female. Ventral 91 69 49 Altamaha River, 10 mi. N Raxley, Appling Co., Geor- margin slightly curved. Dorsal margin gia. Female. straight terminating in a broad angle with 79 63 50 As above. Male. the obliquely descending posterior margin. Hinge ligament prominent, running under Anatomy. Discussed by Lea (1863: 413). the umbos and appearing anteriorly. Pos- Habitat. Usually found in sand bars, terior ridge very sharp and angular near where the river is wide and the water is the umbos, gradually becoming less so rather swift, though occasionally found in toward the ventral margin, especially in mud. older individuals. Posterior slope slightly Remarks. Lampsilis dolabraeformis (Lea) convex, usually with a few faint ridges and is known only from the Altamaha River wrinkles. Umbos swollen, much elevated system. It is close to Lampsilis cariosa above the hinge line, located somewhat (Say) which is found throughout the anterior to the middle of the shell, their Atlantic Slope region, but differs from sculpture consisting of four or five rather cariosa by having umbos which are much evenly raised bars which terminate at the higher and a posterior ridge which is con- posterior ridge. Surface of the shell smooth siderably sharper, especially toward the except for a few concentric ridges ante- umbos. riorly. Periostracum usually shiny, yellowish In the Apalachicolan region, L. dolabrae- green, sometimes becoming chestnut brown formis resembles L. excavatus (Lea). The or black in matures. The posterior slope latter does not extend east of the Escambia usually has a concentration of green or River system. It is a smaller species than black rays which extend with less fre- dolabraeformis, the umbos are not as high, quency over the posterior portion of the the posterior ridge is not quite as sharp, disk. especially in females, and the entire sur- Left valve with two compressed pseudo- face has wide green or blackish rays which cardinal teeth, one in front of the other, are often broken into concentric bands of In the are the anterior one higher. Sometimes these green. dolabraeformis rays restricted to the area toward the are broken into three imperfect teeth or posterior ridge and posterior slope, and the rays are are united in one narrow ridge. Hinge line and not broken. L. dolabrae- rounded, with a considerable interdentum fine, sharp, formis also resembles L. binominatus under the umbo before two short lateral Simpson (Johnson, 1967b: 9) of the Upper teeth. Right valve with two opposing Apalachicola River system. The rays are pseudocardinals, the anterior one low and very similar, but in binominatus they cover laminate, the other and higher, chunky, the entire surface. Further, binominatus is serrated; one lateral tooth, trun- shaqily a much smaller species, which does not cated behind. Beak cavities and deep wide, have the high umbos or sharp posterior with dorsal muscle scars under the deep ridge of dolabraeformis. pseudocardinal teeth. Anterior adductor Simpson (1914, 1: 46) reports dolabrae- muscle scars deep, posterior ones distinct, formis from the Ogeechee and Savannah but not impressed. Pallial line distinct. River systems, but the specimens in the Nacre white, bluish white, or pinkish, United States National Museum are all usuallv iridescent. Lampsilis cariosa (Say). 386 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

Range. Southern Atlantic re- Soc, 4: 118, pi. 18, fig. 46 (Ohio River; Slope: USNM stricted to the Altamaha River system, figured holotype 84509). Lea, 1834, Obs. Unio, 1: 128. Georgia. Unio lenis Conrad 1838, Monography Unionidae, no. 11, back cover. Conrad, 1840, op. tit., no. Specimens Examined 12, p. 106, pi. 58, fig. 2 (upper part of White River, Illinois, possible type ANSP 42307, River System Altamaha [smaller than figured specimen]). Unio canadensis Lea 1857, Proc. Acad. Nat. Sci. River Drainage. Georgia: Ocmulgee Phila., 9: 85 (St. Lawrence River, near Mon- Pulaski Co. Ocmulgee River, Hawkinsville, treal [Quebec], Canada). Lea, 1860, Jour. Ocmulgee River, 1.5 mi. S Jacksonville; Acad. Nat. Sci. Phila., ser. 2, 4: 268, pi. 44, Gum Swamp Creek, 1 mi. N McRae; fig. 148; figured holotype USNM 84503. Lea, 1860, Obs. Unio, 7: 86. Ocmulgee River, 1 mi. S Lumber City; all Unio latissimus Conch. Telfair Co. Sowerby 1868, Iconica, 16, Unio, pi. 66 fig. 337 (United States, Oeonee River Drainage. Georgia: Sowerby coll. not in BMNH [lost]). Oconee River, 2.5 mi. N Glenwood; Och- Lampsilis ventricosa lurida Simpson 1914, Cat. walkee Creek, 2 mi. E Glenwood; Oconee Naiades, 1: 41 (throughout the St. Lawrence River, 8 mi. SW Soperton; all Wheeler Co. drainage; holotype USNM [not located]). Lampsilis ventricosa cohongownta Ortmann 1912, Altamaha River Drainage. Georgia: Nautilus, 26: 53 (no type selected. Lecto- Altamaha River, 7 mi. N Hazlehurst, Jeff type, selected by Parodiz (1967: 28) Carnegie Davis Co. Altamaha 10 mi. River, N Mus. 61.3999, from Potomac River, Hancock, Baxley; Altamaha River, 10 mi. NE Sur- Washington Co., Maryland), pi. 21: 1; allotype rency; both Appling Co. Altamaha River, 61.4000, pi. 21:2. "Riverside Park," 4 mi. N Jesup, Wayne Lampsilis ventricosa winnchagoensis Baker 1928, Co. Altamaha River, 3 mi. NW Everett Bull. Wisconsin Geol. Nat. Hist. Survey, no. 70(2), p. 291, pi. 94, figs. 1-4 (Winnebago City, Glynn Co. Altamaha River, Fort Lake, near Oshkosh [Winnebago Co.], Wiscon- Barrington; Altamaha River, Hopeton, near sin; holotype MZUM 209219). Darien; both Mcintosh Co. Lampsilis ventricosa perglohosa Baker 1928, Bull. Wisconsin Geol. Nat. Hist. Survey, no. 70(2), Lampsilis (Lampsilis) ovata (Say) p. 285, pi. 93, figs. 1-4 (Lake Pepin, near Plate 21: 1-2 Lake City [Wabasha Co.], Wisconsin; holotype MZUM 209195). Unio ovatus Say 1817, Nicholson's Encyclopedia, Lampsilis ovata (Say). Ortmann, 1919, Mem. 2 [no pagination], pi. 2, Fig. 7 (Ohio River Carnegie Mus., 8: 297, pi. 17, figs. 8, 9; pi. and its tributary streams; type, ANSP [lost]). 18, figs. 1-3. Neotype Senckenberg Mus. 4338 [not seen] Lampsilis ovata ventricosa (Barnes). Ortmann, selected, but not figured, by Haas, 1930, 1919, Mem. Mus., 8: Senckenbergiana, 12: 328. Carnegie 301, pi. 18, fig. 4 pi. 19, figs. 1-3. Clarke and Berg, 1959, Lampsilis cardium Rafinesque 1820, Ann. Gen. Cornell Univ. Exp. Sta. Mem., no. 367, p. 55, des Sci. Physiques (Rruxelles), 5: 298, pi. 30, figs. 47, 48. figs. 16-19; ([Ohio River]: supposed type ANSP 20210 teste Proc. Acad. Vanatta, 1915, Description. Shell usually medium, though Nat. Sci. Phila., 66: 5511). reaching 140 mm in length. Outline of Unio ventricosus Barnes 1823, Ainer. Tour. Sci., male elliptical or subelliptical, of female 6: 267, pi. 13, fig. 14 (The Wisconsan [Wis- consin River] near Prairie du Chien [Crawford subovate, rather short and high. Valves Co., Wisconsin] type, Lyceum of Natural somewhat to considerably inflated, thin of New York History [destroyed by fire]). when young, becoming thick with age, in- Unio occidens Lea 1829, Trans. Amer. Philos. equilateral. Anterior end regularly rounded; Soc, 3: 435, pi. 10, fig. 16 (Ohio; figured posterior end more rounded and holotype USNM 84866). Lea, 1834, Obs. broadly in the truncated Unio, 1: 49. somewhat pointed male, Unio suhovatus Lea 1831, Trans. Amer. Philos. in the female. Ventral margin slightly Atlantic Slope Unionidae • Johnson 387

short Length Ileiuht Width curved. Dorsal margin and straight, mm nun mm in a broad or terminating angle gentle 115 78 56 Potomac- River, Hancock, curve with the obliquely descending pos- Washington Co., Maryland. terior margin. Hinge ligament prominent, Female. occupying most of the dorsal margin. Pos- 111 73 47 South Branch, Potomac River, South Branch, Hamp- terior ridge variable; it may be rounded shire Co., West Virginia. and defined, often poorly though becoming Male. Carnegie Mus. distinct toward the umbos, and in some 95 60 41 Potomac River, Hancock, the posterior ridge becomes very Washington Co., Maryland. sharp toward the umbos. Posterior slope Lectotype of L. v. cohon- goronta Ortmann. Male. varies from gently convex to almost flat, or even concave toward the umbos. Umbos 87 52 57 As above. Allotype. Fe- male. moderately swollen, slightly raised above the located somewhat anterior hinge line, Anatomy. Discussed by Ortmann (1911: to the middle of the their shell, sculpture 351). The glochidia are subelliptical, with rather coarse consisting of four or five a straight line hinge, without hooks, and bars, of which the second and third have measure 0.25 mm in length and 0.29 mm a slight tendency to fall into two loops, in height. Coker, et al. (1921: 153) re- with a small sinus in the middle, while the port the following fish as hosts: bluegill, first, fourth, and fifth bars are indistinct. Lepomis macrochirus Rafinesque; white Surface of the shell generally smooth ex- crappie, Pomoxis annularis Rafinesque; salmoides cept for growth ridges. Periostracum largemouth bass, Micropterus smallmouth bass, M. dolo- usually light or dark yellowish green to (Lacepede); mieui dolomieui olive brown. Dark green or blackish rays Lacepede; yellow perch, Perca flave.scens (Mitchill); and yellow which are straight and continuous, finer or pikeperch, Stizostedion vitreum (Mitchill). broader, often cover the entire surface or Breeding season. According to Ortmann only part of it, but some rays are almost (1919: 298, 303, 308) this species is brady- On the the always present. posterior slope tictic. is less smooth and periostracum usually Habitat. Lives in rivers and creeks on darker. gravel, sand, or even mud bottoms; also Left valve with two slightly compressed occurs in lakes. teeth, one in front of the pseudocardinal Remarks. Lampsilis ovata (Say) is a the anterior one somewhat other, triangu- species of the Interior Basin and St. Law- lar, the hinder one lower, located directly rence River system which was accidently under the umbos, both teeth quite chunky. introduced into the Shenandoah River in Hinge line rounded with a narrow inter- 1889 and in other parts of the Potomac dentum under the umbos; two short lateral River system in 1894, while in the larval on of bass other fishes teeth. Right valve with two opposing stage, species and that were at those times from pseudocardinals, the anterior one low and transplanted the west. On the Atlantic Slope ovata laminate, the other higher, quite chunky, can be confused only with Lampsilis and serrated; one lateral tooth, sharply cariosa (Say), with which it is now found truncated behind. Beak cavities moder- in the lower Potomac River system. L. ately deep, with deep dorsal muscle scars. ovata generally has a sharper posterior Anterior adductor muscle scars well im- ridge, especially near the umbos, tends to Pallial line pressed, posterior ones less so. be inflated, has a dull olivaceous tint to distinct. Nacre silvery or bluish white, the shell, is inclined to be rayed over the sometimes suffused with pink or purple. entire surface, and has rather heavy chunky 388 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

pseudocardinals. L. cariosa always has a Marshall (1930: 19) indicated that cariosa poorly defined posterior ridge, is not much was being replaced by ovata. This is con- inflated, usually has a distinctive bright firmed by collections made in the Potomac waxy yellow glossy periostracum. Rays River in 1965 by S. L. H. Fuller. when present are usually confined to the Range. Interior Basin: Mississippi and posterior slope, and the pseudocardinals Ohio drainages. St. Lawrence drainage are less chunky than those of ovata. from Lake Superior to the Ottawa River The present synonymy of ovata may not and Lake Champlain. Hudson Bay drain- the Northern Atlantic restricted to be the last word spoken on the subject ( age. Slope: author is not yet prepared to comment on the Potomac River system, Maryland (in-

is . the validity of Unio satur Lea, which troduced ) found in the Western Gulf Coastal region ) , Specimens but it is an attempt to use modern system- Examined atic concepts. Ortmann (1919: 298, 303) Potomac River System clearly states that L. ovata occurs in large Potomac River West Vir- rivers and gradually gives way to ovata Drainage. ventricosa in smaller rivers and streams, ginia: South Branch, Potomac River, Rom- South Branch, that numerous intergrades occur in inter- ney (Carnegie Mus.); Potomac South Branch = mediate areas and, further, that this is a River, [Depot French both general phenomenon and occurs in widely Station] (Carnegie Mus.); Co. 1 mi. above separated streams. Van der Schalie (1938: Hampshire Cacapon River, confluence with Potomac 70) discusses a similar change from ovata River, Morgan Co. Potomac ventricosa, the river form, to the ovata (USNM). Maryland: River, Co. canadensis, the lake form, which also inter- Hancock, Washington (Carnegie Shenandoah River, grade. Goodrich and van der Schalie Mus.). Virginia: Harp- ers Co. (1944: 315) again note that, "as one pro- Ferry, Jefferson (Carnegie Mus.). Potomac Paton gresses into the headwaters the sharp pos- Maryland: River, Island, 0.75 mi. Point of Rocks, Frederick Co. terior ridge of the true ovata is seen to W Potomac 2.5 mi. round off and we pass gradually to the River, WSW Dickerson; Potomac Harrison Island, 5 mi. more common form of the species in Indi- River, above Great Falls Potomac ana, known as LampsUis ovata ventricosa (USNM); ." River, Great Falls (USNM); Chesapeake ( Barnes ) Cvancara's ( 1963 ) work ap- and Ohio Great Falls all pears to indicate that ovata and ventricosa Canal, (USNM); Co. are ecophenotypes. Montgomery Ortmann (1912: 53) proposed cohongo- ochracea ronta as a variety of LampsUis ventricosa, LampsUis (LampsUis) (Say) Plate 21: 5 "on account of the small size, the shorter 4, and higher outline, and somewhat more Mytilus fluviatilis Gmelin 1791, Systema Naturae, ed. 3359 dulci- distinct posterior ridge." He found no 13, 1, pi. 6, p. (Europae, aquis bus [erroneous], type [probably based on difference in the anatomy or glochidia from Lister's figure only]) refers to: Lister, 1685, L. ventricosa. Synopsis Methodicae Conehyliorum, pi. 157, When Ortmann wrote in 1912, LampsUis fig. 12 (Virginia; figured specimen Oxford Univ. ovata was found on the Atlantic Slope only Mus. [probably lost] teste Dance, 1966, Shell in the Great Allegheny Valley, an area Collecting, p. 292). Conrad (1836, Monog- from which cariosa has never been re- raphy Unionidae, no. 4, p. 37) considered Lister's to ochracea ported. Since that time ovata has spread figure represent LampsUis (Say), but the figure is sufficiently ambiguous throughout the Potomac River, having to have subsequently allowed Isaac Lea to reached Great Falls, 1915 Maryland, by claim successfully, for over fifty years, that (Marshall, 1917: 40). Writing again, M. fluviatilis Gmelin was Anodonta cataracta Atlantic Slope Umomdak • Johnson 389

Say. Since Lister makes no reference to disk, becoming roughened with perios- dentition or lack of it, his figure cannot be tracum posteriorly. Periostracum usually positively identified. Gmelin's name is certainly subshiny, brownish olive, brownish, green- a nomen dubium. ish reddish or Dull, Unio ochraceus Say 1817, Nicholson's Encyclo- yellow, yellow, yellow. pedia, 2 [no pagination], pi. 3 fig. 8 (Delaware rather fine, greenish rays sometimes found The and Schuylkill Rivers; type ANSP [lost]). over the entire surface of the shell, espe- was restricted to Schuylkill River, type locality cially in immatures. mar Philadelphia, Pennsylvania, and a neotype Left valve with two ser- MCZ 178838 was selected by Johnson, 1947, compressed, rated one in front Occ. Papers on Moll., 1: 153, pi. 20, fig. 2. pseudocardinal teeth, Lampsilis rosea Rafinesque 1820, Ann. Gen. des of the other, the anterior one somewhat Sci. 5: 229 (Hudson Physiques (Bruxelles), triangular, the hinder one lower, inclined River [New York]; type not in ANSP [lost]). to be vestigial. Hinge line rather long and Unio wsaceus Conrad 1849, Proc. Acad. Nat. Sci. narrow before two short, curved lateral Phila., 4: 153 (Savannah River, Georgia, J. H. Couper [loaned]). Conrad, 1850, Jour. Acad. teeth. Right valve with two triangular, Nat. Sci. Phila., ser. 2, 1: 275, pi. 37, fig. 5; narrow, rjarallel pseudocardinals separated figured holotype MCZ 178779, purchased from by a deep narrow pit, the more anterior J. H. Couper. tooth inclined to be vestigial; one lateral Lampsilis ochracca (Say), Simpson, 1914, Cat. tooth. Beak cavities with Naiades, 1: 49. Ortmann, 1919, Mem. Car- shallow, deep negie Mus., 8: 318, pi. 20, figs. 6, 7. Johnson, dorsal muscle scars. Anterior adductor 1947, Occ. Papers on Moll., 1: 150, pi. 20, muscle scars well impressed, posterior ones 1-2. Clarke and 1959, Cornell figs. Berg, less so. Pallial line not very distinct. Nacre Univ. Exp. Sta. Mem., no. 367, p. 57, figs. white, bluish white, or usually iri- 55, 56. Athearn and Clarke, 1962, Natl. Mus. pinkish, descent. Canada, Bull. no. 183, p. 30, pi. 4, figs. 3, 4.

Length Height Width Description. Shell usually small, seldom mm mm mm 70 in Outline of male exceeding mm length. 73 44 27 Savannah River, Georgia. elliptical and somewhat elongate; of female Holotype of U. rosaceus Conrad. Male. subovate, rather short and high. Valves 50 31 19 near inequilateral, subinflated, rather thin, Schuylkill River, Philadelphia, Pennsylvania. though strong, sometimes translucent. An- Neotype. Male. terior end regularly rounded; posterior end 54 38 24 Halfway Pond, Plymouth, more rounded and somewhat broadly Plymouth Co., Massachu- pointed in the male, truncated in the fe- setts. Female. male. Ventral margin usually straight or Anatomi/. Figured by Lea (1838: pi. 15, slightly curved, sometimes concave in fig. 44) and described (1863: 455) also by females. Dorsal margin straight or slightly Reardon (1929: 1, pi. 1, figs. 1-10). The curved, terminating in a broad angle with host fish is unknown, but since ochrocea the obliquely descending posterior margin. is generally restricted to the lower regions Hinge ligament prominent, running under of streams or bodies of water directly con- the umbos and appearing anteriorly. Pos- nected with the ocean, it may, like Ano- terior ridge rounded and generally poorly donta implicata Say, parasitize a migratory defined. Posterior slope slightly convex, fish. with a few faint and usually ridges Breeding season. Lea (1863: 455) found wrinkles. Umbos moderately swollen, this species gravid in the autumn. I found slightly raised above the hinge line, located gravid females on May 3, 1943 in Plym- somewhat anterior to the middle of the outh, Massachusetts. It is probably brady- shell, their sculpture consisting of four or tictic. five wavy recurved ridges. Surface of the Habitat. Lives in sand or mud in ponds, shell generally smooth in the region of the canals, and the lower portions of rivers to- 390 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

ward the tidal region in areas where the Waccamaw; drainage canal, beside Lake current is not strong. Waccamaw, 1 mi. NNW Dupree Landing; Remarks. In the Atlantic Slope region, both Columbus Co. Soutli Carolina: Lampsilis ochracea (Say) has been con- Waccamaw River, Wachasaw Landing, 2 fused with L. cariosa (Say), but ochracea mi. W Murrells Inlet, Georgetown Co. has a thinner, smaller shell and, unlike Pamlico River System cariosa which is rarely rayed, ochracea is often rayed over the entire surface. It has Tar River Drainage. North Carolina: a rougher, duller, more greenish perios- Tar River, Old Sparta, 3.5 mi. W Pinetops, tracum. The hinges are quite different. Edgecombe Co. Tar River, Bruce, 9 mi. The pseudocardinals of cariosa are pyra- NW Greenville, Pitt Co. midal and an interdentum is present, but Roanoke River System in ochracea the pseudocardinals are lamel- late, almost parallel to the hinge line, and Roanoke River Drainage. North Caro- there is no interdentum. lina: Roanoke River, Weldon, Halifax Lampsilis ochracea may also be mis- Co. (USNM). taken for immature Lampsilis splendida (Lea), but only in that the shells of both James River System be over the entire surface. The may rayed Appomattox River Drainage. Virginia: of ochracea are and the shell is rays finer, Petersburg, Dinwiddie Co. (USMN). more whereas that of yellowish, splendida James River Drainage. Virginia: James is more brownish, less delicate, and more River (MCZ and USNM). inflated. Simpson (1914: 49) gives the range of York River System this species as from "New England to the Mattaponi River Drainage. Virginia: The Ogeechee River, Georgia." specimen Mantapike Creek, Mantapike, King and in the United States National Museum on Queen Co. (USNM). which he based the Ogeechee record from York River Drainage. Virginia: York Le Conte to be a cariosa. appears young River (MZUM). Unfortunately, in none of the major mu- seums are there any precisely labeled spec- Potomac River System imens of ochracea from the Savannah River, Potomac River Drainage. Virginia: Po- the apparent southern limit of its distribu- tomac River, Great Falls, Fairfax Co. tion. Conrad (1836: 37) reports ochracea District of Columbia: Potomac River. Ana- as occurring in, "most of the tide waters costia River. Virginia: Canal, Alexandria north of Savannah River." (MZUM); Little Hunting Creek, near Mt. Atlantic Savannah River Range. Slope: Vernon MZUM both Fairfax Co. ( ) ; system, Georgia, north to River Herbert, Nova Scotia (Athearn and Clarke, 1962: Lampsilis (Lampsilis) radiata radiata 30). (Gmelin) Plate 22: 1-3 Specimens Examined Mya radiata Gmelin 1791, Systema Naturae, ed. 13, 1, pt. 6, p. 3220 (Malabariae [erroneous]) Savannah River System refers to: Martini and Chemnitz, 1782, Conch.

7 ( [source of Cmelin's Savannah River Drainage. Georgia: Cabinet, 6, pi. 2, fig. colln. Savannah River (MCZ and USNM). locality]; figured specimen Spengler Copenhagen, [lost] teste Haas, 1913, Vidensk. Waccamaw River System Meddr. Dansk naturh. Foren. [Copenhagen], 65: 59); also Lister, 1685, Synopsis Methodicae Waeeamaw River North Drainage. Conchyliorum, pi. 152, fig. 7 (Virginia; figured Carolina: Lake Waccamaw, [town of] Lake specimen, Oxford Univ. Mus. [probably lost] Atlantic Slope Unionioae • Johnson 391

teste, Dance, 1906, Shell Collecting, p. 292). 1 .amarck's type examined and identified as The figure in Martini and Chemnitz is here Elliptio complanatus Solander [sic] teste John- selected as the type figure, and the type lo- son, 1953, Nautilus, 66: 95. cality, following Lister and Simpson (1914, Lampsilis radiata (Gmelin). Simpson, 1914, Cat. Cat. Naiades, 1: 64), is here restricted to Naiades, 1: 64. Ortmann, 1919, Mem. Car-

Potomac River, District of Columbia ( approxi- negie Mus., 8: 292, pi. 17, figs. 6, 7. mately opposite, Fairfax Co., Virginia). Lampsilis radiata oneidensis Baker 1916, Nautilus, [//i/'o hiteola Lamarck 1819, Hist. Nat. des Ani- 30: 74, pi. 2 (Oneida Lake, Central New York; maux sans Vertebres, 6: 79 (la riviere Susque- syntypes, New York State College of Forestry, hana [Susquehanna River, Pennsylvania] et Syracuse, New York; ANSP 114854; MZUM). celle Mohancks [Mohawk River, New York]; Unio virginca Frierson 1927, Check List North measured holotype Paris Museum, figured by American Naiades, p. 72, error for Unio vir- Wheeler, 1963, Nautilus, 77: 58, pi. on p. 59, giniana Lamarck 1819, non Lamarck. figs. 1-2. Refigured by Johnson, 1969, Nautilus, Lampsilis radiata radiata (Gmelin). Clarke and 83: 54, fig. 11, the type locality was restricted Berg, 1959, Cornell Univ. Exp. Sta. Mem. no. York to the Susquehanna River, Columbia, Co., 367, p. 58, fig. 53. Athearn and Clarke, 1962, Pennsylvania ) . Natl. Mus. Canada, Bull. no. 183, p. 31, pi. 4, St. [7/i/'o lincata 'Valenciennes' Bory de Vincent figs. 5, 6. 2 of 1827, Encyclopedic Methodique, atlas, Lampsilis radiata var. conspicua (Lea). Simpson, 5 explanation of pis., p. 151, pi. 248, fig. (no 1914, Cat. Naiades, 1: 66. locality, type [presumed lost]). [Named from figure published in 1797.] Description. Shell large, often reaching [//i/'o tenebrosus Conrad New Fresh Water 1834, over 100 mm in length. Outline subellipti- Shells United States, p. 42, pi. 7, fig. 1 (Rap- cal or subovate, moderately elongated, pahannock [sic] River, near Falmouth, Stafford a half times as in Poulson distinctly over one and long Co., Virginia; tvpe' presumably colln., ANSP [lost]). as high. Valves generally subinflated, [7/i/o melinus Conrad 1838, Monography Unioni- though occasionally quite inflated, and no. 101, 1 (Salina Lake dae, 11, p. pi. 56, fig. solid. Anterior end regularly rounded; jdos- [Onondaga Co.], New York; type, not in ANSP, terior end more broadly rounded, and in presumed lost). females often somewhat in the Unio boydianus Lea 1840, Proc. Amer. Philos. expanded not so. Soc, p. 286 (Oak Orchard Creek, Orleans Co., joostbasal region, though uniformly New York). Lea, 1843, Trans. Amer. Philos. Ventral margin generally regularly curved, 8: Soc, 216, pi. 16, fig. 32; figured holotype sometimes straight, roughly parallel to the USNM 86126). Lea, 1942, Obs. Unio, 3: 54. almost straight dorsal margin which forms Unio rosaceus De Kay 1843, Zool. New York, a blunt angle with the posterior margin in Moll., pt. 5, p. 192, pi. 39, figs. 355, 356; pi. and becomes almost 40, fig. 357 (Seneca Lake [North Western], immatures, impercep- New York; lectotype, here selected, USNM tible with age. Hinge ligament prominent. 678303, formerly New York State Cabinet 391, Posterior ridge feeble or wanting. Posterior specimen figured on pi. 39, fig. 356). slojie broad and undistinguished. Umbos Mija oblongata Wood 1856, Index Testaceologicus, rather sharp, but not full or high, their ed. Hanley, p. 199, Supplement, pi. 1, fig. 2 of six to ten double- (North America, type [lost]). sculpture consisting with a distinct Unio elongata S. G. Goodrich 1859, Illustrated looped bars, re-entering Natural History, 2: 523, text fig. (No locality, angle in the middle. The posterior looj) is not type [probably saved].) slightly angular and indistinct on the jdos- Unio Reeve 1856, Conch. Iconica, obliquiradiatiis terior slope. Surface roughened by close 16, Unio, pi. 29, figs. 151 (Hab?, Mus. concentric wrinkles of periostracum, gen- Cuming, type [not in BMNH, lost]). or brownish with Unio conspicuus Lea 1872, Proc. Acad. Nat. Sci. erally yellowish green, or blackish both nar- Phila., 24: 156 (Yadkin River, Salisbury, dark greenish rays, [Rowan Co.], North Carolina). Lea, 1874, row and wide, over the entire surface. Acad. Nat. Sci. ser. 8: Jour. Phila., 2, 34, pi. Left valve with two stumrjy pseudo- 11, fig. 31; figured holotype USNM 85056. cardinal teeth, both rather triangular and Lea, 1874, Obs. Unio, 13: 38. crenulate. No interdentum. Two long, Unio virginiana Simpson, 1900, Proc. United teeth. valve States Natl. Mus., 22: 536, non Lamarck 1819. slightly curved lateral Right 392 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

with one rather high, triangular pseudo- under splendida on p. 394. Lampsilis r. cardinal, and a vestigial one before it; one radiata, an Atlantic Slope species, most lateral tooth. Beak cavities very shallow, closely resembles Lampsilis radiata sili- with a few dorsal muscle scars. Anterior quoidea (Barnes) which is found through- and posterior adductor muscle scars and out the Interior Basin. Clarke and Berg pallial line all distinct. Nacre generally (1959: 58-62, 68-70) have shown that in white, bluish white, sometimes tinted with the Lower St. Lawrence drainage of New pink or salmon, sometimes quite pink or York where these two otherwise quite salmon. distinct "species" commingle, they inter-

Length Height Width grade completely. mm mm mm Typically, L. r. radiata is easily dis- 138 86 46 Lake 12 mi. Michie, NNE tinguished from L. r. siliquoidea. The Durham, Durham Co., former is more compressed; the difference North Carolina. Male. between the male and female shell is much 112 64 39 Yadkin River, near Salis- less marked; the is bury, Rowan Co., North periostracum always Carolina. Holotype of U. rough and not shiny like that of siliquoidea. conspicuus Lea. Male. It has broader rays and often has a reddish 99 52 29 Susquehanna River, Co- or salmon nacre. In siliquoidea the nacre is York Penn- lumbia, Co., always bluish white. sylvania. Male. Ortmann (1919: 296) suggested for the 84 49 27 Potomac River, Washing- type locality of r. radiata Saratoga Lake ton, District of Columbia. Male. in New York, "should there not be any 62 39 20 As above. Female. earlier record," and was probably unaware that Simpson (1914, 1: 65) had already Lea ( 1838, pi. 15, Anatomy. Figured by selected Virginia as the type locality. to Ortmann 1911: figs. 48, 49). According ( Ortmann and Walker (1922: 61) have the of L. radiata radiata 349) anatomy shown that Unio luteolus Lamarck is not "in all essential with that agrees respects" recognizable on the basis of the original of L. r. The siliqaoidca (Barnes). glochidia description. Wheeler (1963: 58) has tried and measure are suboval, without hooks, to resurrect this name on the basis of the 0.22 to 0.23 mm in and 0.27 to 0.28 length identification of the type. Lamarck origi- mm in The host fish is unknown. height. nally gave two localities, the Susquehanna Breeding season. Ortmann (1919: 293) and the Mohawk rivers. The former has indicates that the season breeding begins only L. r. radiata, and the latter, which is in and ends the August following August. now in the region of hybridization, prob- Habitat. Lives in rivers and lakes of all ably did not have L. r. siliquoidea in it in or sizes, usually gravel sand, occasionally before the completion of the Erie Canal, but ascends in mud. Prefers tidewaters, some years after Lamarck's specimen was some of the larger rivers, such as the collected. Therefore, to avoid further con- the Susquehanna River, Pennsylvania, and fusion the type locality was restricted to Hudson River, New York, for a consider- the Susquehanna. See: synonymy above. able distance. Simpson (1914, 1: 66) treats Unio con- Remarks. In the Southern Atlantic Slope spicuus Lea as a variety of Lampsilis r. region Lampsilis radiata radiata (Gmelin) radiata, stating that "there are intermedi- is replaced by Lampsilis splendida (Lea) ates, which seem to fully connect the two." below the Pedee River system. But the I am in agreement with him. The main two species do not much resemble one difference in the southern specimens seems another, as splendida has a sharp posterior to be a general tendency to grow large ridge and r. radiata docs not. See: Remarks more consistently than elsewhere. There Atlantic Slope Unionidae • Johnson 393

does seem to be a hiatus for this species Unio regularis Sowerby 1866, Conch. Iconica, 16, between the Neuse River, North Carolina, Unio, pi. 34, fig. 181, non Lea 1841. Lampsilis splendida Lea. Simpson, 1914, Cat. and the Potomac River, Virginia. Ortmann Naiades, 1 : 50. (1919: 296) calls attention to the fact that while this species is abundant in the tide- Description. Shell large, often exceeding water region of the Potomac River, it does 110 mm in length. Outline obovate and not go up the river into the mountains west elongated, the female proportionally higher of the Blue Ridge, and is rare or absent on than the male. Valves considerably in- the Piedmont Plateau. flated, subsolid. Anterior end rounded; of the Range. Southern Atlantic Slope: dis- posterior end male broader and very continuous, Pedee River system, South bluntly pointed; the female even more Carolina, Cape Fear and Neuse river broadly rounded. Ventral margin usually systems, both North Carolina. Northern slightly curved, or slightly arcuate in males Atlantic Slope: Potomac River, Maryland, when they are swollen behind the middle of the north to the St. Lawrence River system, base. Dorsal margin straight, some- westward to Lake Ontario. times forming a distinct angle with the obliquely descending posterior margin or Specimens Examined merging gradually into it. Hinge ligament prominent, under the umbos and Pedee River System running appearing anteriorly. Posterior ridge rather Yadkin River Drainage. North Caro- sharp, generally with a second ridge above lina: [Yadkin River], Salem, Forsyth Co. it. Posterior slope considerably wrinkled. Yadkin (MZUM). River, near Salisbury, Umbos very high and full, their sculpture Rowan Co. (MZUM). consisting of a number of strong, nearly straight bars. Surface of the disk some- Cape Fear River System times smooth, but the periostracum is Cape Fear River Drainage. North Caro- usually concentrically wrinkled, especially lina: Greenfield Pond, Wilmington, New posteriorly and toward the margins. Perios- Hanover Co. (USNM). tracum yellowish green to brownish with the entire surface covered with numerous Neuse River System wide and narrow green rays. Old shells Neuse River Drainage. North Carolina: sometimes dark brown with the rays ob- Lake Michie [an impoundment on Flat scured. River], 12 mi. NNE Durham Center, Dur- Left valve with two compressed pseudo- ham Co. (75). cardinal teeth slightly anterior of the umbos, one in front of the other, the an- Potomac River System terior one somewhat triangular; the hinder one Potomae River Drainage. Virginia; Po- considerably lower. Hinge line very tomac River, Great Falls, Fairfax Co. narrow, before two short, curved, lateral District of Columbia: Potomac River. Vir- teeth. Right valve with two triangular, ginia: Mount Vernon, Fairfax Co. narrow, parallel pseudocardinals separated by a deep narrow pit, the more anterior Lampsilis (Lampsilis) splendida (Lea) tooth inclined to be vestigial; the hinder Plate 22: 5 4, tooth inclined to be chunky in old speci- Unio splendidus Lea 1838, Trans. Amer. Philos. mens. One rather high lateral tooth present. Soc, 6: 70, pi. 19, fig. 61 (Altamaha River, near Beak cavities deep and wide, with dorsal Darien [Mcintosh Altamaha Co.]; [River], muscle scars. Anterior adductor muscle Liberty [now Long] Co.; both Georgia; figured scars well ones less so. holotvpe USNM 84893). Lea, 1838, Obs. Unio, impressed, posterior 2: 70. Pallial line distinct. Nacre white, orange, 394 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

violet, sometimes bluish, but more inclined to be pinkish and iridescent.

Length Height Width

112 Atlantic Slope Unionidae • Johnson 395

SELECTED BIBLIOGRAPHY Anodonta fragilis Lamarck. Natl. Mus. Canada, Bull. 199. Cont. Zool., 1963: 15-27. The references were selected to include those Clench, W. J. 1962. Collecting freshwater mol- which are cited in the as well as briefly text, lusks in south central Georgia. Shells and those which are especially relevant to the area their neighbors, 12: 1, 7. [A popular publi- under study. cation, discontinued in 1964.]

Adams, C. C. 1901. Baseleveling and its faunal Clench, W. J., and K. J. Boss. 1957. Freshwater significance, with illustrations from the Mollusca from James River, Virginia. Nauti- Southeastern United States. Amer. Natur., lus, 80: 99-101.

35: 839-851. Clench, W. J., and R. D. Turner. 1956. Fresh-

Adams, G. I. 1926. In Adams, G. I., et al., water mollusks of Alabama, Georgia and Geology of Alabama. Geol. Survey Alabama, Florida from the Escambia to the Suwannee Spec. Rept. no. 14: 25-27. River. Bull. Florida State Mus., 1: 97-239. Alt, D. 1968. Pattern of Post-Miocene eustatic Conrad, T. A. 1834 (May). New fresh water fluctuation of sea level. Paleogeog., Paleoclim., shells of the United States with colored Paleoecol, 5: 87-94. illustrations, and a monograph of the genus Alt, D., and H. K. Brooks. 1965. Age of Anculotus of Say; also a synopsis of the Florida marine terraces. Tour. Geol., 73: American naiades. Philadelphia, Penn., pp. 406-411. 1-76.

Athearn, H. D. 1964. Three new unionids from . 1835 (October), ibid. Appendix, pp. 1-8, Alabama and Florida and a note on Lamp- pi. 9 (col.). silis joncsi. Nautilus, 77: 134-139. . 1836-40. Monography of the family of Athearn, H. D., and A. H. Clarke, Jr. 1962. Unionidae, or naiades Lamarck, ( Fresh

The freshwater mussels of Nova Scotia. Natl. water bivalve shells ) of North America. Phila- Mus. Canada, Bull. 183, Cont. Zool., 1960- delphia, Penn., pp. i-v, 1-118 [pp. 13-16 61: 11-41. never printed]. Published in thirteen parts.

Bailey, J. L., Jr. 1940. Wilmington, N. G, . 1846. Notices of fresh water shells, etc., Records. Nautilus, 54: 69. Elliptio fisheri- of Rockbridge Co., Virginia. Amer. Jour. (11111.1 Lea [= Elliptio htnccolata (Lea)], Sci., ser. 2, 1: 405-407. Lake Greenfield, Wilmington, N. G, is the 1853. A synopsis of the family of naiades only unionid mentioned. of North America, with notes, and a table of Ball, G H. 1922. Variation in freshwater mus- some of the genera and sub-genera of the sels. Ecology, 3: 93-121. family, according to their geographical dis-

Bates, J. M. 1966. A new species of Canmculina tribution, and descriptions of genera and sub- (Unionidae: Pelecypoda) from the Savannah genera. Proc. Acad. Nat. Sci. Phila., 6: 243- River, South Carolina. Occ. Pap. Mus. Zool., 269. Univ. Mich., no. 646: 1-9. Cooke, C. W. 1925. Physical geography of

Boss, K. J., and W. J. Clench. 1967. Notes on Georgia: The coastal plain. Geol. Survey Pleurohema collina (Conrad) from the James Georgia, Bull., no. 42: 19-54.

River, Virginia. Occ. Pap. Moll., Mus. Comp. . 1930. Pleistocene seashores. Jour. Wash. Zool., 3: 45-52. Acad. Sci., 20: 389-395. R. E. 1896. A revision and of Call, synonymy . 1936. Geology of the coastal plain of the parvus group of Unionidae. Proc. Indiana South Carolina. Geol. Survey, Washington, Acad. Sci., 1895: 109-119. D. G, Bull., no. 867: 1-12. M. R. 1896. modifications Campbell, Drainage . 1943. Geology of the coastal plain of and their interpretation. Jour. Geol., 4: 567- Georgia. Geol. Survey, Washington, D. G, 581. Bull., no. 941: 103-112. Cherry, R. N. 1961. Chemical quality of water 1945. Geology of Florida. Florida Geol. of Georgia streams, 1957-58. Georgia State Survey, Geol. Bull., no. 29: 1-339. Division Conservation. Geol. Bull, of Survey Cvancara, A. M. 1963. Clines in three species no. 69: 1-100. of Lampsilis (Pelecypoda: Unionidae). Mala- Clarke, A. H., Jr., and G O. Berg. 1959. The cologia, 1: 215-225. freshwater mussels of central New York. Dawley, G 1965. Checklist of freshwater mol- Cornell Univ. Agr. Expt. Sta., Mem. no. 367: lusks of North Carolina. Sterkiana, no. 19:

1-79. 35-39. An uncritical list; most of the identi-

Clarke, A. H., Jr., and A. M. Rick. 1963. Sup- fications appear to be based on whatever plementary records of Unionacea from Nova appeared on the labels of specimens examined Scotia with a discussion of the identity of or are from the literature. 6 396 Bulletin Museum of Comparative Zoology, Vol. 140, No.

of com- Emery, K. O, 1967. The Atlantic continental — . 1947b. The audiorship EHiptio 62: 36. margin of the United States dining the past planatus. Nautilus,

of — . 1956. The of Naiades (Mollusea: 70 million years. Geol. Assn. Canada, types in the Museum of Special Paper 4, Geol. of the Atlantic Region, Unionidae) Comparative 115: 101- pp. 53-70. Zoologv. Bull. Mus. Comp. Zool., Flint, R. F. 1957. Glacial and Pleistocene ge- 142. Anodonta ology. New York, pp. i-xiii, 1-553. . 1965. A hitherto overlooked Frierson, L. S. 1912. Notes on Anodonta (Mollusea: Unionidae) from the Gulf drain- couperiana and A. gibbosa. Nautilus, 25: age of Florida. Breviora, Mus. Comp. Zool., 129-130. no. 213: 1-7. redi- of all the mollusks . 1915. Lasmigona subviridis Conrad, . 1967a. Illustrations vivus. Nautilus, 29: 57-59. described by Berlin Hart and Samuel Hart check -. 1927. A classified and annotated Wright. Occ. Pap. Moll., 3: 1-35. naiades. list of the North American Waco, . 1967b. Additions to the unionid fauna Texas: Baylor Univ. Press, pp. 1-111, errata of the Gulf drainage of Alabama, Georgia sheet. Includes a reference to almost all of and Florida (Mollusea: Bivalvia). Breviora, the taxa, within its scope, descrihed before Mus. Comp. Zool., no. 270: 1-21. 1927. an . 1967c. Carunculina pulla (Conrad), Gibbes, L. R. 1848. Mollusea [of South Carolina]. overlooked Atlantic drainage unionid. Nauti- on the of In Tuomey, M., Report geology lus, 80: 127-131. South Carolina. pp. xix-xxii. unionid Appendix, . 1968. EHiptio nigella, overlooked Goodrich, C. 1930. Unio spinosa Lea. Nautilus, from Apalaehicola River system. Nautilus, 43: 140. 82: 22-24.

. of the 1939. Certain mollusks Ogeechee . 1969a. Further additions to the unionid River, Georgia. Nautilus, 52: 129-131. fauna of the Gulf drainage of Alabama, Nothing on Unionidae, but refers to van der Georgia and Florida. Nautilus, 83: 34-35. Schalie's expedition of 1937. 1969lx Illustrations of Lamarck's types Goodrich, C., and H. van der Schalie. 1944. of North American Unionidae mostly in the A revision of the Mollusea of Indiana. Amer. Paris Museum. Nautilus, 83: 52-61. 32: 257-326. Midland Nat., Keith, A. 1925. Physical geography of Georgia: Gould, A. A. 1848. Descriptions of shells found The highland. Geol. Survey Georgia, Bull., in Connecticut, collected and named by the no. 42: 93-132. Amer. late Rev. J. H. Linsley. Jour. Sci., Kurten, B. 1966. Holarctic land connexions in ser. 6: 233-236. 2, the early Tertiary. Comm. Biol. Soc. Sci. 1926. Further Grier, N. M., and J. F. Mueller. Fennica, 29(5): 1-5. of and station in studies in correlation shape Laessle, A. M. 1968. Relationship of sand pine Bull. Free fresh water mussels. Wagner scrub to former shore lines. Quart. Jour. Instit. 1: 11-28. Sci., Philadelphia, Florida Acad. Sci., 30: 270-286. 1969a. Unionacea. In Das Haas, F. Superfamilia LaForce, L. 1925. Physical geography of Geor- Tierreich. Berlin. Lief. 88: I-X, 1-663. gia: The central upland. Geol. Survey In . 1969b. Unionacea. Superfamily Georgia, Bull., no. 42: 57-92. Treatise on Invertebrate Part N. Paleontology, Lea, I. 1834-74. Observations on the genus N411-N471. Mollusea 6 (Bivalvia), pp. Unio. Philadelphia, Penn., 1-13. A reprint R. Campbell. 1894. Hayes, C. W., and M. of Lea's papers from various journals. They of the southern Geomorphology Appalachians. are repaged, but the plate and figure num- Natl. 6: 63-126. Geographic Mag., bers are the same as those in the original Heard, W. II. 1966. Population sexuality in journals. Amer. Malacol. Ann. Anodonta. Union, Rept. structure . 1838. [Notes on the anatomical for 1966: 31-33. of Naiades]. Trans. Amer. Philos. Soc, 6: 1967. Pleistocene Hoyt, J. H., and J. R. Hails. 48-57; also 1838, Obs. Unio, 2: 48-57. shoreline sediments in coastal Georgia: Depo- . 1854. Rectification of Mr. T. A. Conrad's sition and modification. Science, 155: 1541- of the of naiades of North 1543. "Synopsis family America," published in the "Proceedings of Johnson, R. I. 1946. Anodonta implicata Say. the of Natural Sciences of Phila- Occ. Papers Moll., Mus. Comp. Zool., I: 109- Academy Proc. Acad. Nat. 116. delphia," February, 1853. and Sci. Phila., 7: 236-249. . 1947a. Lampsilis cariosa Say Lamp-

. 1858. of the silis ochracea Say. Occ. Pap. Moll., Mus. Descriptions embryonic forms of of Unionidae. Comp. Zool., 1 : 145-156. thirty-eight species Atlantic Slope Unionidae • Johnson 39"

Jour. Acad. Nat. Sci. Phila., ser. 2, 4: 43-50, Myers, G. S. 1938. Fresh-water fishes and West also Obs. 6: pi. 5; 1858, Unio, 43-50. Indian zoogeography. Smithsonian Rept., 1863. Descriptions of the soft parts of [for] 1937: 339-364. one hundred and forty-three species and Oaks, R. Q., and N. K. Coch. 1963. Pleistocene some embryonic forms of Unionidae of the sea levels, southeastern Virginia. Science, United States. Jour. Acad. Nat. Sci. Phila., 140: 979-983. ser. 2, 5: 401-456; also 1863, Obs. Unio, Ortmann, A. E. 1906. The crawfishes of the 10: 37-92. state of Pennsylvania. Mem. Carnegie Mus., Linsley, J. H. 1845. Catalogue of the shells of 2: 343-524.

Connecticut. Amer. Jour. Sci., 48: 271-286. . 1909. The breeding season of Unionidae MacNeil, F. S. 1950. Pleistocene shore lines in in Pennsylvania. Nautilus, 22: 91-95, 99- Florida and Georgia. Geol. Survey, Washing- 103. ton, D. C, Prof. Paper 221-F, pp. 95-106. . 1910. A new system of the Unionidae. Marshall, W. B. 1890. Beaks of Unionidae in- Nautilus, 23: 114-120. habiting the vicinity of Albany, New York. . 1911. A monograph of the najades of Bull. New York State Mus., 2: 169-189. Pennsylvania. Parts 1 and 2. Mem. Carnegie

. 4: 279-347. 1917. Lampsilis oentricosa cohongoronta Mus., in the Potomac River. Nautilus, 31: 40-41. . 1912a. Notes upon the families and of the . 1918. Lampsilis ventricosa cohongoronta genera najades. Ann. Carnegie Mus., 8: 222-365. in the Potomac Valley. Nautilus, 32: 51-53.

. 1912b. . 1930. Lampsilis ventricosus cohongoronta The geological origin of the fresh-water of in the Potomac River. Nautilus, 44: 19-21. fauna Pennsylvania. Penn. Matteson, M. P. 1948a. The taxonomic and Topog. Geol. Survey, 1910-12: 130-149.

. 1913a. distributional history of the freshwater mussel The Alleghenian Divide and its influence the freshwater fauna. Proc. EUiptio complanatus ( Dillwyn, 1817). Nauti- upon Amer. lus, 61: 127-132; 62: 13-17. Philos. Soc, 52: 287-390. [This is a classic with which I . have noted 1948b. Life history of EUiptio com- study, only a few minor planatns (Dillwyn, 1817). Amer. Midland disagreements.] Nat., 40: 690-723. . 1913b. Studies in najades. Nautilus, 27: 88-91. . 1955. Studies on the natural history of -. 1914. Studies in the Unionidae. Amer. Midland Nat., 53: najades. Nautilus, 28: 126-145. 41-47, 65-69, 129-131.

. 1915. Studies in Mayr, E., E. G. Lixsley, and R. L. Usinger. najades. Nautilus, 29: 63-67. 1953. Methods and Principles of Systematic . 1919. A on the Zoology. New York, McGraw-Hill, pp. 1- monograph naiades of 336. Pennsylvania. Part 3. Systematic account of the and Mem. Mazyck, W. G. 1913. Catalog of Mollusca of genera species. Carnegie Mus., 8: 1-384. South Carolina. Contributions from the i-xiv,

. 1920. Correlation of Charleston Museum No. 2: 1-39. [Includes shape and station in freshwater mussels. an uncritical list of Unionidae based on the Proc. Amer. Philos. Soc. 19: 269-312. literature and on previous identifications of specimens in the museum.] . 1923. The anatomy and taxonomy of certain Unionidae and Anodontinae from the McCallie, S. W. 1925. Physical geography of Gulf 36: 73-84. Georgia. Introduction: The state as a whole. drainage. Nautilus, -. 1923-1924. Notes on the Geol. Survey Georgia, Bull., no. 42: 1-17. anatomy and taxonomy of certain from the Modell, H. 1942. Das natiirliche system der Lampsilinae Gulf 37: Najaden. 1. Archiv fur Molluskenkunde, 74: drainage. Nautilus, 56-60, 99-104, 161-191. 137-144. Ortmann, A. E., and B. Walker. 1922. On the . 1949. Das natiirliche system der Najaden. nomenclature of certain North American 2. Archiv fur Molluskenkunde, 78: 29-48. naiades. Occ. Pap. Mus. Zool. Univ. Mich., -. 1964. Das natiirliche system der Najaden. no. 112: 1-75. 3. Archiv fur Molluskenkunde, 93: 71-126. Parodiz, J. J. 1967. Types of North American [Authorized English translation of part 1 by Unionidae in the collection of the Carnegie Mu- D. H. Stansbery and U. Soehngen, 1964, seum. Sterkiana, no. 28: 21-30. [A lectotype Sterkiana, no. 14: 1-18.] is selected for L. v. cohongoronta Ortmann G. E. 1961. Murray, Geology of the Atlantic male: C. M. 61.3999 and female: C. M. and Gulf Coastal Province of North America. 61.4000 as allotype, p. 28. The author was New York. 1-523. pp. unaware that I had published a list of the 398 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

types of B. H. and S. H. Wright (Johnson, and North and South Carolina. Nautilus, 62: 1967a) a few months previous to his paper.] 121-126. [Only two Unionidae are noted: ochraceus and com- Parodiz, J. J., and A. A. Bonetto. 1963. Taxon- Lampsilis (Say) Elliptio omy and zoogeographic relationships of the planatus quadrilaterus (Lea)=E. complan- ata both from Lake South American naiades (Pelecypoda: Unio- ( Lightfoot ) , Waccamaw, aacea and Mutelacea). Malaeologia, 1: 179- North Carolina.] 213. -. 1967. Valid zoological names of the Portland Proc. U. S. Natl. Patrick, R., J. Cairns, Jr., and S. Roback. 1967. catalogue. Mus., An ecosystematic study of the fauna and flora 121, 1-15. of the Savannah River. Proc. Acad. Nat. Sci. Russell, R. J. 1957. Instability of sea-level. Phila., 118: 109-407. [Unionidae, pp. 345- American Scientist 45: 414-430. 346.] [The following species of Unionidae are Say, Thomas. 1830-34. American conchology, or listed from several localities: (1) Elliptio descriptions of the shells of North America, il- hopetonensis (Lea)=E. complanata (Light- lustrated by colored figures. New Harmony, foot), (2) E. incrassatus (Lea) = E. con- Indiana. 68 col. pis. with letterpress. Pub- garaea (Lea), (3) E. lanceolata (Lea), (4) lished in 7 parts. [Reprinted by: Binney, Uniomerus obesus (Lea) = U. tetralasmus William G. 1858. The complete writings of (Say), (5) Anodonta couperiana Lea, (6) Thomas Say on the conchology of die United A. hallenbecki Lea = A. cataracta Say, (7) States. New York. 71 col. and 4 plain pis.]

A. imbecilis Say, (8) Lampsilis eariosa (Say), SCHNABLE, J. E., AND H. G. GOODELL. 1968. (9) L. splendida (Lea), (10) Villosa ogee- Pleistocene-Recent stratigraphy, evolution and cheensis (Conrad) = V. delumbis (Conrad).] development of the Apalachicola Coast, Pilsbry, H. A. 1894. Critical list of mollusks Florida. Geol. Soc. Amer., Special paper 112, collected in the Potomac Valley. Proc. Acad. pp. 1-72. Nat. Sci. Phila., 46: 11-30. Sellards, E. H., and H. Gunter. 1918. Geology Rafinesque, C. S. 1831. Continuation of a between the Apalachicola and Ochlockonee monograph of the hivalve shells of the river Rivers in Florida. Geology between the Ohio, and other rivers of the western states. Choctawhatchee and Apalachicola Rivers in Philadelphia, Penn. pp. 1-7. [This, and the Florida. lOth-llth Ann. Rept. Florida Geol. papers from the various journals, were re- Surv., pp. 9-56, pp. 77-102. and printed by: Binney, William C, George Sickel, J. B. 1969. A survey of the mussel popu- of W. Tryon. 1864. The complete writings lations ( Unionidae ) and Protozoa of the Constantine Smaltz Rafinesque on recent and Altamaha River with references to their use fossil conchology. Philadelphia, Penn. pp. in monitoring environmental changes, pp. 2 1-96, 7, 3 pis.] [Mimeographed abstract of a master's thesis, Raulerson, L., and W. D. Burbanck. 1962. The Emory University, Atlanta, Georgia.] life cycle and ecology of Elliptio hopetonen- Simpson, C. T. 1892. Notes on the Unionidae sis Lea. Assoc. Southeast. Biol., 9: 39. [An of Florida and the southeastern states. Proc. abstract of a master's thesis, "The ecology U. S. Natl. Mus., 15: 405-436. [Many of of a small Georgia mountain stream, with the figures are line drawings of the types.]

special emphasis on the mussel, Elliptio . 1900. Synopsis of the naiades, or pearly hopetonensis Lea," by Claire Lynn Raulerson freshwater mussels. Proc. U. S. Natl. Mus.,

. 22: 501-1044. ( 1960, Emory University, Atlanta, Georgia ) I misidentified these specimens; they are E. -. 1914. A descriptive catalogue of the naiades or freshwater mussels. Detroit, complanata ( Lightfoot ) .] pearly Ravenel, E. 1874. Catalogue of the recent and Michigan. Parts 1-3, pp. xi, 1-1540 fossil shells in the cabinet of Edmund Simpson, G. B. 1884. Anatomy and physiology Ravenel. [Edited by L. R. Gibbes] Charles- of Anodonta fluviatilis. Thirty-fifth Ann. ton, South Carolina, pp. 1-67. [Not seen. Rept. New York State Mus. Nat. Hist., pp. Includes a nomen nudum, Alasmidonta 169-191. coarctata Ravenel (rivers in the upper part Stephenson, L. W. 1912. In W. B. Clark, et al., of South Carolina), p. 58.] The Coastal Plain of North Carolina. North Reardon, L. 1929. A contribution to our knowl- Carolina Geol. Econ. Surv., 3, The Quater- edge of the anatomy of the fresh-water nary Formations, pp. 266-290. mussels of the District of Columbia. Proc. Swain, G. F., et al. 1899. Papers on the water- U. S. Natl. Mus., 75: 1-12. power in North Carolina. North Carolina Rehder, H. A. 1949. Some land and freshwater Geol. Survey: Bull., no. 8. mollusks from the coastal region of Virginia Thomas, G. J., and D. C. Scott. 1965. Note Atlantic Slope Unionidae • Johnson 399

on EUiptio spinosa in Georgia. Nautilus, 79: INDEX TO RELEVANT TAXA 66-67. Principal references in boldface type. Tomkins, I. R. 1955. Elliptic spinosus in the Altamaha River. 68: 132-133. Nautilus, abbevillensis Lea, Unio, 315 E. G. 1915. Vanatta, Rafinesque's types on aberrans Lea, Unio, 317 Unio. Proc. Acad. Nat. Sci. Phila., 67: ablatus Lea, Unio, 327 549-559. Acglia Swainson, 382 van der Schalie, H. 1938. The naiad fauna aequatus Lea, Unio, 326 of the Huron River, in southeastern Michi- alabumensis Lea, Margaritana, 366 Misc. gan. Pub. Mus. Zool., Univ. Michigan, Alasmidonta Say, 346, 348 no. 40: 1-83. Alasminota Ortmann, 343 . 1940. The naiad fauna of the Chipola Alasmodonta Say, 348 River, in northwestern Florida. Lvoydia, 3: ampins Lea, Unio, 318 191-208. [Published by the Lyoyd Library, angulata (Lea), Gonidea, 265 Cincinnati, Ohio.] angustatus Lea, Unio, 333 . 1945. The value of mussel distribution annulatus Sowerby, Anodon, 367 in tracing stream confluence. Pap. Michigan Anodonta Lamarck, 356 Acad. Sci. Arts Letters, 20: 355-373. anodontoides (Lea), Lampsilis, 271

. 1966. Hermaphroditism among North anthonyi Lea, Unio, 312 American freshwater mussels. Malacologia, Anodontinae (Swainson) Ortmann, 343 5: 77-78. aquilus Lea, Unio, 326 van der Schalie, H., and A. van der Schalie. arctata (Conrad), EUiptio (EUiptio), 271, 274, 1950. The mussels of the Mississippi River. 276, 331 American Midland Nat., 44: 448—466. arctatus Conrad, Unio, 331

Vaughn, T. W. 1910. A contribution to the arcula (Lea), Alasmidonta (Alasmidonta), 27'4, geologic history of the Floridian plateau. 278, 352 Carnegie Inst., Washington, Pub. no. 133: arcula Lea, Margaritana, 352 99-185. areolatus Swainson, Anodon, 367 Walker, R. 1905. List of shells from north- arkansasensis Lea, Anodonta, 367 western Florida. Unionidae from Moccasin atropurpureum Rafinesque, Alasmidon, 353 Creek, a tributary of the Econfine River. attentuata Rafinesque, Obliquaria, 380 Nautilus, 18: 136. attenuatus Lea, Unio, 380 aurata 315 . 1910. The distribution of Margaritana Rafinesque, Unio, australis 272 margaritifcra ( Linn. ) in North America. Proc. Simpson, Lampsilis, Mai. Soc. London, 9: 126-145. averellii B. H. Wright, Unio, 373 -. 1918. A synopsis of the classification of baldwinensis the fresh-water Mollusca of North America. Lea, Unio, 316 barrattii 326 Misc. Pub. Mus. Zool. Univ. Michigan, no. Lea, Unio, 6: 1-213. basalis Lea, Unio, 318 beaverensis Lea, Unio, 318 . 1919. Notes on North American naiades. binominatus Simpson, Lampsilis, 273, 385 II. Unio tenerus Ravenel and Lampsilis bisselianus Lea, Unio, 340 modioUformis (Lea). Occ. Pap. Mus. Zool. 339 Univ. Michigan, no. 74: 1-8. blandingianus Lea, Unio, boydianus Lea, Unio, 391 Walter, W. M. 1956. Mollusks of the upper boijkiniana (Lea), Amblema, 271 Neuse River Rasin, North Carolina. Jour. brimleyi S. H. Wright, Unio, 301 Elisha Mitchell Sci. Soc, 72: 262-274. buddianus Lea, Unio, 339 Walter, W. M., and R. A. Parker. 1957. EUip- Bullella Simpson, 348 tio complanatus roanokensis in the Neuse burkei Walker, Quincuncina, 273 River. 71: 60-64. Nautilus, hurkensis Lea, Unio, 316 C. K. 1930. Sand and re- Wentworth, gravel burtchianus S. H. Wright, Unio, 327 sources of the coastal plain of Virginia. Vir- buxcus Lea, Unio, 308 ginia Geol. Survey, Rull. no. 32: 1-146. Wheeler, H. E. 1935. Timothy Abbott Conrad, calceola (Lea), Alasmidonta, 279, 349 widi particular reference to his work in camptodon Say, Unio, 339 Alabama one hundred years ago. Rull. Amer. canadensis Lea, Unio, 386 Paleont., 23(77): 1-157. Canthijria Swainson, 303 Wheeler, M. J. 1963. Type of Unio luteolus cardium Rafinesque, Lampsilis, 386 Lamarck 1819. Nautilus, 77: 58-61. carinifera Lamarck, Unio, 314 400 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

cariosa (Say), Lampsilis (Lampsilis), 274, 275, cuspidatus Lea, L/n/o, 327 276, 278, 279, 382 cuvierianus Lea, L/n/o, 315 cariosus Say, Undo, 382 cygneus Linnaeus, Mytilus, 356, 362 CaruncuUna Baker, 369 cylindraceus Frierson, Elliptio, 327 castas Lea, Unio, 301 cataracta Say, Anodonta (Pyganodon) cataracta, danielsii B. H. Wright, C/ra'o, 305 272, 274, 276, 278, 279, 286, 356 dariensis Lea, Anodonta, 357 catawbensis Lea, Unio, 315 dariensis (Lea), Etf/pf/o (Elliptio), 274, 277, 278, charlottensis Lea, Unio, 344 310 chathamensis Lea, Unio, 317 dariensis Lea, l/n/o, 310 318 chipolancnsis (Walker), Elliptio, 273 datus, Lea, L/n/o, declivis 339 choctawensis Athearn, Villosa, 273 Say, l/n/o, dear Lea, Unio, 340 decoratus Lea, l/n/o, 344 Decurambis cirratus Lea, L/n/o, 319 Rafinesque, 353 delumbis 375 cistelliformis Lea, l/n/o, 317 Conrad, l/n/o, delumbis claibornensis (Lea), Lampsilis, 271, 382 (Conrad), V///osa, 274, 276, 277, 375 diazensis S. H. coarctata Lamarck, [/mo, 314 Wright, l/n/o, 327 cohongoronata Ortmann, Lampsilis ventricosa 386 differtus Lea, l/n/o, 318 dilatata 305 collina Conrad, Pleurobcma (Lexingtonia), 274, (Rafinesque), Elliptio, 270, 279, 280, 277, 300 dilatatus Rafinesque, Unio, 305 discus 305 collinus Conrad, Unio, 300 Sowerby, Unio, B. H. 327 columbensis Lea, l/nio, 340 dispalans Wright, l/n/o, dispar Lea, L/n/o, 373 complanata ( Lightfoot ) , Elliptio (Elliptio), 272, dissimilis 319 274, 278, 279, 286, 314 Lea, l/n/o, complanata (Lightfoot), Mija, 314 dolabraeformis (Lea), Lampsilis (Lampsilis), 274, 384 compressa (Lea), Lasmigona, 280, 343, 345 278, 384 compressa Lea, Symphynota, 343 dolabraeformis Lea, L/n/o, dolearis 357 concavus Lea, l/n/o, 376 Lea, Anodonta, doliaris 357 confertus Lea, l/rao, 325, 329 Lea, Anodonta, 319 congaraea (Lea), Elliptio (Elliptio), 274, 276, dooleyensis Lea, l/n/o, dorsatus 277, 278, 308 Lea, L/n/o, 308 downiei congaraeus Lea, l/n/o, 308 (Lea), Elliptio (Elliptio) crassidens, 267, 307 conica Rafinesque, Pleurobcma, 299 273, downiei 307 conspicuus Lea, l/n/o, 391, 394 Lea, L/n/o, Anodonta, 365 constricta (Conrad), V/Z/o.sa, 274, 277, 278, 280, dunlapiana Lea, duttonianus 285, 378 Lea, l/n/o, 333 constrictus Conrad, Unio lienosus, 378 edentula Say, Alasmidonta, 367 contiguus Lea, Unio, 376 cdentulus Say, Strophitus, 368 contractus Lea, l/n/o, 316 elect rinus Reeve, l/n/o, 340 contrarius Conrad, l/n/o, 382 £///p//o Refinesque, 303, 304 corneas Lea, C/ra'o, 319 clongata Goodrich, l/n/o, 391 corrugata DeKay, Alasmidon, 354 emmonsii Lea, l/m'o, 333 Corunculina Simpson, 370 errans Lea, l/n/o, 315 coruscus Gould, Unio, 326 cscambia Clench and Turner, Fusconaia. 272 corvus Lea, l/n/o, 308 Eunjria Rafinesque, 305 costata Rafinesque, Ala-smidonta, 243 Anodonta exactus Lea, l/n/o, 316 coupcriana Lea, ( Utterbackia ) , 272, 274, exacutus Lea, l/n/o, 334 275, 276, 365 excavatus (Lea), Lampsilis, 271, 385 cowperiana Lea, Anodonta, 365 excultus Conrad, l/n/o, 339 crassidens (Lamarck), Elliptio (Elliptio) cras- excurvata DeKay, Anodon, 356 sidens, 267, 271, 277, 305 exiguus Lea, l/n/o, 373 crassidens- var. b. Lamarck, C/mo, 305 extensus Lea, C/rwo, 333 crocatus Lea, l/n/o, 382 cuneata Rafinesque, Pleurobcma, 299 fishcrianus Kuester, l/n/o, 280 cuncalus Barnes, Unio, 305 fishcrianus Lea, C/rao, 333, 334 Cunicula Swainson, 305 fisheropsis De Gregorio, l/n/o arctior, 334 cuprea Rafinesque, Obliquaria, 333 floridensis (Lea), Lampsilis anodontoides, 270 curatus Lea, l/n/o, 317 fluviatilis Gmelin, Mytilus, 388 curvatus Lea, L'mo, 318 fluviatilis Green, l/n/o, 315 Atlantic Slope Unionidae • Johnson 401

folliculatus Lea, Unto, 333 implicata Say, Anodonta, 360 forbesianus Lea, Unto, 308 implicate Say, Anodonta (Pyganodon), 274, 275, fragilis Lamarck, Anodonta cataracta, 266, 279, 279, 360 357 incerta Lea, Anodonta, 363 fraterna (Lea), EUiptio (Elliptic), 272, 274, 276, incrassatus Lea, C/nio, 305 277, 312 indefinitus Lea, C/nio, 317 fraternus Lea, Unto, 312 indifinilus Lea, C/nio, 317 fryanus B. H. Wright, C/niO', 327 ineptus Lea, C/nio, 339 fuliginosus Lea, C/nio, 315 infulgens Lea, C/nio, 319 fulvus Lea, C/nio, 308 inf areata (Conrad), Quincuncina, 272 fumatus Lea, C/nio, 316 infuscus Lea, C/nio, 318 /(/.«•« Baker, Utterbackia imbecUlis, 363 insolidus Lea, C/nio, 344 fuscatus Lea, C/nio, 326 insulins Lea, C/nio, 315 Fusconaia Bafinesque, 300 inusitatis Lea, C/nio, 324 invenustus Lea, C/nio, 319 gastonensis Lea, C/nio, 317 irwinensis Lea, C/nio, 318 geddingsianus Lea, C/nio, 326 geminus Lea, C/nio, 315 jamesianus Lea, C/nio, 340 genthii Lea, C/nio, 378 jayensis Lea, EUiptio, 272, 335 genuinus Lea, C/nio, 376 jejunus Lea, C/nio, 315 geometricus Lea, C/nio, 339 jewettii Lea, C/nio, 340 georgina Lamarck, C/nio, 314 /onesi van der Schalie, Lampsilis, 273 gesnerii Lea, Anodonta, 356 Jugosus Simpson, 348 gesnerii Lea, C/nio, 319 lacustris Lea, Anodonta, 356 gibbesianus Lea, C/nio, 308 lacustris Baker, Strophitus rugosus, 367 gibbosa Say, Anodonta (Pyganodon), 27'4, 278, 359 Lampsilinae (Ihering) Ortmann, 369 glabrata Lamarck, C/nio, 314 Lampsilis Bafinesque, 382 glabratus Sowerby, C/nio, 349 lanceolata globosa Lea, Anodonta, 356 (Lea), EUiptio (EUiptio), 272, 274, 276, 278, 279, 281, 333 gracilenius Lea, C/nio, 332 lanecolatus Lea, C/nio, 333 gracilior Lea, C/nio, 373 lapillus Say, C/nio, 372 grandis Say, Anodonta, 271, 279, 360 Lasmigona Bafinesque, 343 griffithianus Lea, C/nio, 315 Lastcna Bafinesque, 362 haddletoni Athearn, Lampsilis, 373 latissima Bafinesque, Ligumia recta, 380 hallenbeckii Lea, Anodonta, 356, 358 latissimus Sowerby, C/nio, 386 hallenbcckii Lea, C/nio, 316 /a/ns Lea, C/nio, 316 hartwrightii B. H. Wright, C/nio, 310 lazarus Lea, C/nio, 331 hazelhurstianus Lea, C/nio, 333 lazarus Sowerby, C/nio, 317 hefe

margaritifera (Linnaeus), Margaritifera, 265, 266, ouara Valenciennes, Unio, 382 279, 280 ovatus Frierson, Strophitus undulatus, 367 marginata (Say), Alasmidonta, 266, 279, 280, 281, ouarus Say, Unio, 382, 386 355 306 marginata Say, Anodonta, 356 pachijodon Pilsbry, EUiptio, 'Ravenel' 319 masoni (Conrad), Plcurobema (Lexingtonia), 274, palliatus Simpson, Unio, 382 277, 278, 301 pallida Rafinesque, Lampsilis, 340 masoni Conrad, Unio, 301 paludicolor Conrad, Unio, 339 maywebbae B. H. Wright, EUiptio, 327 paludicolus Gould, Unio, 367 mccordi Athearn, Alasmidonta, 270 papyracea Anthony, Anodon, 339 mcmichaeli Clench and Turner, EUiptio, 312, 313 paralellus Conrad, Unio, 371 mecklcnbergensis Lea, Unio, 317 /wufl (Barnes), Carunculina, 271, 276, 277, 370 mediocris Lea, Unio, 317 parvus Barnes, Unio, 370 melinus Conrad, Unio, 391 patrickae Bates, Carunculina, 367 merceri Lea, L/»/o, 327 pavonia Lea, Anodonta, pawensis 340 nierus Lea, C7nio, 301 Lea, Unio, 366 micans Lea, C/n/o, 326 peggyae Johnson, Anodonfa, 272, 362, 364, 271 Micromya Agassiz, 371 penicillatus (Lea), Medionidus, 367 modioliformis Lea, I7mo, 373 pensylvanica Lamarck, Anodonta, 367 Monodonta Say, 348 pepinensis Baker, Strophitus rugosus, percoarctatus 315 monroensis Lea, (7n?o, 310 Lea, Unio, perglobosa Baker, Lampsilis ventricosa, 386 moussonianus Lea, t/nio, 308 332 Mutelacea, 264 perlatus Lea, Unio, 317 mytiloides Rafinesque, Pleurobema, 299 perlucens Lea, Unio, perplicata 271 mytiloides Swainson, t/rao, 348, 349 (Conrad), Amblema, 270, perstriatus Lea, Unio, 332 wesuto (Say), Ligumia, 274, 275, 279, 281, 380 pertcnuis Lea, Unio, 344 nasutus Say, (7mo, 380 planilatcris Conrad, Unio, 308 nasutilus Lea, C7»n'o, 334 plantii Lea, Unio, 340 nasutulus Lea, C/rao, 334 Platynaias Walker, 343 naviculoides Lea, Unio, 333 Pleurobema Rafinesque, 299 neglecta (Lea), Lasmigona, 343 polymorphic B. H. Wright, Unio, 305 neisleri (Lea), Amblema, 273 porrectus Conrad, Unio, 339 neusensis Lea, t/m'o, 316 postellii Lea, Unio, 316 newtonensis Lea, Anodonta, 361 Pressodonta Simpson, 349 nige/Z« (Lea), EUiptio, 273 prevostianus Lea, Unio, 373 n/gra Rafinesque, (7m'o, 305 productus Conrad, Unio, 333 nigrinus Lea, Unio, 373 Prolasmidonta Ortmann, 346 /lo/anj B. H. Wright, L/m'o, 327 protensus Lea, Unio, 317 northamptonensis Lea, t/m'o, 317 provancheriana Pilsbry, Unio (Arconaia), 319 nubilus Lea, Unio, 318 proximus Lea, Unio, 376 Pscudodontoidcus Frierson, 366 obesus Lea, (7nio, 339 Pterosyna Rafinesque, 343 obfuscus Lea, (7m'o, 373 pulla (Conrad), Camnculina, 21A, 276, 277, 370 ohlatus Lea, (7nfo, 327 pullatis Lea, Unio, 326 obliquiradiatus Sowerby, C/n/o, 391 pullatus Lea, Unio, 326 oblongata Wood, t/nio, 391 370 oblongus Lea, (7?i/o, 391 pullus Conrad, Unio, obnubilis Lea, Unio, 326, 329 pumilus Lea, Unio, 301 occidens Lea, C/nio, 386 purpurascens Lamarck, Unio, 305, 314 occultus Lea, Unio, 326 purpureus Say, Unio, 314 ochracea (Say), Lampsilis (Lampsilis), 21 A, 275, 276, purus Lea, Unio, 316 388 279, pusillus Lea, U/iio, 308 ochraccus 389 Say, Unio, Pyganodon Crosse and Fischer, 356 ocmulgeensis Lea, t/nio, 327 pygmaeus Lea, Unio, 344 ogeecheensis Conrad, Unto, 375 pyriforme (Lea), Pleurobema, 272, 276, 277, 302 ohiensis Rafinesque, Anodonta (Lastena), 362 oneidensis Baker, Lampsilis radiata, 391 quadrata (Lea), Lasmigona, 343 opacus Lea, Unio, 326 quadratus Lea, Unio, 316 oratus Conrad, Unio, 382 quadrilaterus Lea, Unio, 317 ouato (Say), Lampsilis (Lampsilis), 274, 279, 386 quadriplicate Sowerby, Anodon, 367 Atlantic Slope Unionidae • Johnson 403

radiata (Gmelin), Lampsilis (Lampsilis) radiata, s/;/no.sy; (Lea), EZZtpfto (Canthyria), 21 A, 278, 303 21A, 275, 277, 278, 279, 390 spinosus Lea, C/n/o, 303 radiata Gmelin, Mya, 390 spissus Lea, C/n/o, 307 radiata Swainson, Unio, 348 splcndida (Lea), Lampsilis (Lampsilis), 274, 276, radiata Swainson, Uniopsis, 349 277, 278, 393 radiatus (Conrad), Anodontoides, 271, 366, 368 splendidus Lea, C/n/o, 393 radiolus Lea, Unio, 327 st pallidas Lea, C/n/o, 340 raensis Lea, Unio, 316 s(piameus Lea, C/n/o, 317 raleighensis Lea, C/n/o, 317 stagnalis Conrad, C/n/o, 373 rarisulcata Lamarck, t/nto, 314 striatulus Lea, C/n/o, 301 ratus Lea, C/n/o, 318 strigosus (Lea), Elliptio, 328, 332 raveneli Conrad, Unio, 325 strigosus Lea, C/n/o, 331 ravencliana (Lea), Alasmidonta, 355 strodeanum (Wright), Plcurobema, 272 recto Lamarck, Undo, 380 Strophitus Rafinesque, 366 regularis Sowerby, C/n/o, 393 strumosus Lea, C/m'o, 309 rhombica Anthony, Alasmodon, 367 subangulata (Lea), Lampsilis, 272 rhombula Lamarck, Unio, 314 subcroceus Conrad, C/n/o, 340 rtgtYfa Wood, Unio, 315 subcylindraceus Lea, C/nfo, 334 rivicolus Conrad, Unio, 340 subellipsis Lea, C/n/o, 373 rivularis Conrad, C/n/o, 340 subflavus Lea, C/n/o, 316 roanokensis Lea, C/n/o, 315 subinflatus Conrad, C/n/o complanatus, 315 rosaceus Conrad, Unio, 389 sublatus Lea, C/m'o, 326 rosaceus DeKay, Unio, 391 subniger Lea, C/n/o, 316 rosea Rafinesque, Lampsilis, 389 subolivaceus Lea, C/n/o, 319 rostcllum Lea, C/n/o, 334 suborbiculata Say, Anodonta, 271, 362 rostraeformis Lea, C/m'o, 333 subovatus Lea, C/n/o, 386 rostrata Valenciennes, C/m'o, 380 subparallelus Lea, C/n/o, 318 rostrum Lea, C/n/o, 317 subplanus Conrad, C/n/o, 300, 301 roswellensis Lea, C/n/o, 316 subrostrata (Say), Ligumia, 279, 281 rotulata (Wright), Obovaria, 273 subsquamosus Lea, C/n/o, 318 rotundata (Lamarck), Glebida, 271 subvexus (Conrad), Strophitus, 271, 274, 343, 366, rubcllum (Lea), Plcurobema, 270 368 rufusculus Lea, C/n/o, 308 subviriclis (Conrad), Lasmigona (Platynaias), 21 A, Rugifera Simpson, 353 275, 276, 279, 280, 285, 343 rugosus Swainson, Anodon, 367 subviridis Conrad, C/n/o, 343 rugulosa Wood, Mya, 354 succissa (Lea), Fusconaia, 272 sudus 373 rutilans Lea, C/m'o, 373 Lea, C/n/o, sulcidens Lamarck, C/n/o, 315 Sulcularia 349 sagittiformis Lea, C/m'o, 333 Rafinesque, 343, swainsoni 349 salcbrosus Lea, C/n/o, 316 Sowerby, C/n/o, symmetricus C/n/o, 339 salmonia Clessin, Anodon, 367 Lea, Symphynota Simpson, 343 santeensis Lea, C/n/o, 318 satillaensis 307 Lea, C/n/o, tappanianus Lea, C/n/o, 344 snfr/r Lea, C/nio, 388 tenebrosus Conrad, C/n/o, 391 savannahcnsis Lea, C/n/o, 316 tenerus Simpson, C/n/o, 375 sai/// Ward, C/m'o, 339 tennesseensis Frierson, Strophitus undtdatus, 366 scriptum Rafinesque, Alasmodon, 353 tore* Conrad, Anodonta, 356 tores 270 sculptilis Say, Alasmidonta, 349 ( Rafinesque ) , Lampsilis, 367 shaefferiana Lea, Anodonta, 367 tetragona Lea, Anodonta, tetralasmus Say, C/n/o, 276, 339 shepardiana (Lea), Elliptio (Elliptio), 21A, 278, 338 tetralasmus 339 shepardianus Lea, Unio, 338 (Say), Uniomerus, 271, 274, tetricus Lea, C/n/o, 326 siliquoidea (Barnes), Lampsilis radiata, 279, 391 tortivus Lea, C/n/o, 331 similis Lea, C/nio, 326 tortuosus Sowerby, C/n/o, 318 simpsoni B. H. Wright, C/n/o, 327 Toxolasma Rafinesque, 369 singularis B. H. Wright, C/n/o, 327 triangulata (Lea), Alasmidonta (Alasmidonta), 272, sloatiana ( Lea 272 ) , Elliptio, 274, 276, 278, 351 Solcnaia Conrad, 265 triangulata Lea, Margaritana, 351 sordidis Lea, C/n/o, 308 tryonii Lea, Anodonta, 357 spadiceus Lea, C/n/o, 315 tuomeyi Lea, C/n/o, 326, 329 404 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

uhareensis Lea, Unio, 318 villosus Simpson, Lampsilis, 372 unadilla DeKay, Anodon, 367 Dillosus B. H. Wright, Unio, 372 undulata (Say), Alasmidonta (Alasmidonta), 274, violaceus Spengler, Unio, 314 275, 279, 281, 349 virens Lea, L/m'o, 316 undulata Say, Anodonta, 366, 367 virgata Conrad, Anodonta, 367 undulata Say, Monodonta, 348 virginea Frierson, C/rao, 391 undulata Say, Um'o, 349 virginiana Lamarck, Unio, 315 undulatus (Say), Strophitus, 274, 276, 279, 280, 367 virginiana Simpson, Unio, 391 unicolor (Lea), Obovaria, 270 virgulata Lea, Anodonta, 356 Unionacea Thiele, 299 viridans Lea, l/nio, 327 Uniomcrus Conrad, 339 viridicatus Lea, L/n/o, 326 Unkmidae (Fleming) Ortmann, 299 viridiradiatus Lea, (7?n'o, 326 Unioninae (Swainson) Ortmann, 299 viridis Rafinesque, Lasmigona, 343 Unionoidea Thiele, 299 viridis Conrad, C/m'o, 344 Uniopsis Swainson, 348 viridulus Lea, t/?ik>, 333 utriculum (Lea), Pleurobema, 300 Utterhachia Baker, 362 waccamaivensis (Lea), Elliptio (Elliptio), 274, 278, Utterbackia Baker, 362 313 Utterbackiana Frierson, 362 waccamaivensis Lea, £7nio, 313 walkeri (Wright), Medionidus, 272 vanuxemensis (Lea), Villosa, 277, 280, 281, 379 wardiana Lea, Anodonta, 367 varicosa (Lamarck), Alasmidonta (Decurambis), 274, watereensis Lea, t/ra'o, 325 275, 279, 354 websterii B. H. Wright, l/n/o, 310 varicosa Lamarck, Unio, 354 iveldonensis Lea, Unio, 317 vaughanianus Lea, Unio, 375 ivheatleyi Lea, l/m'o, 315 vaughanianus Sowerby, Unio, 380 whiteianus Lea, C/n/o, 326 ventricosus Barnes, Unio, 386 williamsii Lea, Anodonta, 357 verutus Lea, l/rao, 327 winnebagoensis Baker, Lampsilis ventricosa, 386 vestitus 308 Lea, C7n/o, winnebagoensis Baker, Strophitus rugosus, 367 uj'foex (Conrad), Villosa, 271, 274, 276, 373 wrightiana (Walker), Alasmidonta, 273, 350 vibex Conrad, Unio, 373 wrightianus Clench and Turner, Strophitus, 351 vicinus Lea, t/m'o, 315 wrightianus Walker, Strophitus, 348 ViKosc Frierson, 371, 372 villosa (Wright), Villosa, 272, 372 yadkinensis Lea, L/mo, 318

406 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 1

The Southeastern United States

A.M. The solid line indicates the limit of the Appalachian Mountains. The divide is somewhat to the west of it.

is the Fall Line. P.P. Piedmont Plateau. The line thus indicated, ,

C.P. Coastal Plain. The line thus indicated, '--', is the limit of the Brandywine (maximum) interglacial (now

thought to be Upper Miocene) flooding, and is, in part, conjectural.

A. Area of stream capture of the New River and the North Fork of the Roanoke River.

B. Area of confluence of the headwaters of the Alabama-Coosa, Apalachicola and Savannah River systems.

C. Area of suspected stream confluence between Uphauppee Creek of the Alabama-Coosa River system and Uchee Creek of the Apalachicola River system.

D. Area of suspected confluence of the headwaters of the Choctawhatchee River and the Chattahoochee River.

The Principal Drainage Systems

1. ALABAMA-COOSA RIVER SYSTEM 20. EDISTO RIVER SYSTEM 2. Coosa River Drainage 21. COOPER-SANTEE RIVER SYSTEM 3. Tallapoosa River Drainage 22. Saluda River Drainage 23. Broad River Drainage Apalachicolan Region 24. Catawba River Drainage 4. ESCAMBIA RIVER SYSTEM 25. BLACK RIVER SYSTEM

5. YELLOW RIVER SYSTEM 26. PEDEE RIVER SYSTEM

6. CHOCTAWHATCHEE RIVER SYSTEM 27. Yadkin River Drainage 7. APALACHICOLA RIVER SYSTEM 28. WACCAMAW RIVER SYSTEM 8. Chattahoochee River Drainage 29. CAPE FEAR RIVER SYSTEM 9. Flint River Drainage 30. NEUSE RIVER SYSTEM 10. OCHLOCKONEE RIVER SYSTEM 31. PAMLICO RIVER SYSTEM 11. ECONFINA RIVER SYSTEM 32. ROANOKE RIVER SYSTEM 12. SUWANNEE RIVER SYSTEM 33. Dan River Drainage 13. ST. MARYS RIVER SYSTEM 34. Roanoke River Drainage 14. SATILLA RIVER SYSTEM 35. CHOWAN RIVER SYSTEM 36. JAMES RIVER SYSTEM Southern Atlantic Slope Region Northern Atlantic 15. ALTAMAHA RIVER SYSTEM Slope Region 16. Ocmulgee River Drainage 37. YORK RIVER SYSTEM 17. Oconee River Drainage 38. RAPPAHANNOCK RIVER SYSTEM 18. OGEECHEE RIVER SYSTEM 39. POTOMAC RIVER SYSTEM 19. SAVANNAH RIVER SYSTEM 40. Shenandoah River Drainage Atlantic Slope Unionidae • Johnson 407

SOUTHEASTERN UNITED STATES

JO 408 Bulletin Museum of Comparative Zoology, Vol 140, No. 6

PLATE 2

Pleurobema (Lexingfonia) collina (Conrad)

a branch of the James River Lec- Fig. 1. Unio collinus Conrad. North [= Maury] River, [Rockbridge County], Virginia. totype ANSP 41007. Length 46 mm, height 30 mm, width 19 mm (nat. size).

with ammonium chloride After Boss Fig. 2. [James River] USNM 84376. Length 18.8 mm. Specimen sprayed vapor. and Clench (1967, pi. 15, fig. 2).

Pleurobema [Lexingtonia] masoni (Conrad)

ANSP 41333. Fig. 3. Unio masoni Conrad. Savannah River, Augusta [Richmond County], Georgia. Holotype Length 26.6 mm, height 19.3 mm, width 13.3 mm (nat. sire).

234387. 47 28 width 19 Fig. 4. Mill Race, 2 mi. N of Sardis, Burke County, Georgic, MCZ Length mm, height mm, mm (nat. size).

ANSP 41332. Fig. 5. Unio masoni Conrad. Savannah River, Augusta [Richmond County], Georgia. Paratype Length 37 mm, height 26 mm, width 16 mm (nat. size).

40 27 width 17 mm (nat. Fig. 6. Unio casfus Lea. South Carolina. Holotype USNM 84782. Length mm, height mm, size).

North Carolina. Fig. 7. Unio brimleyi Wright. [Walnut Creek of] Neuse River, Raleigh [Wake County], Lectotype USNM 149651. Length 38 mm, height 29 mm, width 15 mm (nat. siz.e).

South Carolina. USNM 85698. Fig. 8. Unio merus Lea. Abbeville District [Savannah River drainage], Holotype Length 38 mm, height 29 mm, width 15 mm (nat. size).

North Carolina. USNM Fig. 9. Unio pumi'/us Lea. Black River [about 10 mi. W of Benson, Johnston County], Holotype 84545. Length 28 mm, height 20 mm, width 13 mm (nat. size).

North Carolina. USNM 84548. Fig. 10. Unio striatums Lea. Roanoke River, Weldon [Halifax County], Holotype Length 39 mm, height 28 mm, width 17 mm (nat. size).

Elliptio (Canthyria) spinosa (Lea)

234055. 59 37 Fig. 11. Altamaha River, 4 mi. NE of Jesup, Wayne County, Georgia. MCZ Length mm, height mm, width 22 mm (approximately 1.2 X)- Atlantic Slope Unionidae • Johnson 409 410 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 3

Elliptio (Elliptio) crassidens crass/dens [Lamarck)

Fig. 1. Unio lehmanii Wright. St. Marys River [Nassau County], Florida. Lectotype USNM 149650. Length 60 mm, height 40 mm, width 24 mm (nat. size).

Fig. 2. Unio polymorphus Wright. Spanish Creek [a tributary of the St. Marys River W of Folkston], Charleton County,

Georgia. Lectotype USNM 152060. Length 75 mm, height 46 mm, width 29 mm (slightly reduced).

Fig. 3. Spanish Creek, Charleton County, Georgia. MCZ 269228. Length 62 mm, height 40 mm, width 30 mm (slightly reduced).

Fig. 4. As above. Length 94 mm, height 57 mm, width 38 mm (slightly reduced). Atlantic Slope Unionidae • Johnson 411 412 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 4

Elliptio (Elliptio) crassidens downiei (Lea)

Fig. 1. Um'o downiei Lea. Buck Lake, a bayou of the Sati 1 1 a River, Wayne [Branfly] County, Georgia. Holotype USNM 84854. Length 84 mm, height 49 mm, width 36 mm (slightly reduced).

Fig. 2. Unio satillaensis Lea. Sati! la River, Camden County, Georgia. Holotype USNM 84855. Length 75 mm, height

45 mm, width 30 mm (slightly reduced).

Fig. 3. Sati II a River, 3 mi. S of Hortense, Brantly County, Georgia. MCZ 237460. Length 27 mm, height 16 mm, width 8 mm (nat. size).

Fig. 4. Unio spissus Lea. Satilla River, Wayne County, Georgia. Holotype USNM 84853. Length 76 mm, height 57 mm, width 40 mm (approximately nat. size). Atlantic Slope Unionidae • Jolinsoii 413 414 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 5

EUiptio (Elliptio) congaraea (Lea)

Fig. 1. Unio congaraeus Lea. Congaree River [Cooper-Santee River system], South Carolina. Holotype USNM 85693.

Length 41 mm, height 25 mm, width 14 mm (slightly reduced).

Fig. 2. Unio (ulvus Lea. South Carolina. Holotype USNM 85679. Length 40 mm, height 21 mm, width 15 mm (slightly reduced).

Fig. 3. Unio vestitus Lea. Ogeechee River [Georgia]. Holotype USNM 85332. Length 39 mm, height 25 mm, width 15

mm (nat. size).

Fig. 4. Unio rorbesianus Lea. Savannah River, Georgia. Holotype USNM 84542. Length 45 mm, height 31 mm, width

29 mm (nat. size).

Fig. 5. Unio leconlianus Lea. Canoochee River, Liberty County, Georgia. Holotype USNM 84852. Length 70 mm,

• ' height 45 mm, width 29 mm (slig-htly reduced). '•'.' •

Fig. 6. Unio dorsatus Lea. Catawba River, North Carolina. Holotype USNM 84494. Length 57 mm, height 40 mm, width 23 mm (nat. size).

Fig. 7. Unio moussonianus Lea. Georgia. Holotype USNM 85168. Length 72 mm, height 49 mm, width 28 mm (slightly reduced).

Fig. 8. Unio corvus Lea. Buckhead Creek, Burke County, Georgia. Holotype USNM 84539. Length 62 mm, height 40 mm, width 25 mm (nat. size). Atlantic Slope Unionidae • Johnson 415 416 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 6

Elliptic [Elliptio] dariensis (Lea)

Fig. 1. House Creek, Bowens Mills, 9 mi. N of Fitzgerald, Ben Hill County, Georgia. MCZ 234237. Length 41 mm, height 23 mm, width 14 mm (nat. size).

Fig. 2. Unio dariensis Lea. [Altamaha River] near Darien [Mcintosh County], Georgia. Holotype USNM 85691. Length

78 mm, height 55 mm, width 28 mm (slightly reduced).

Fig. 3. House Creek, Bowens Mill, 9 mi. N of Fitzgerald, Ben Hill County, Georgia. MCZ 234237. Length 76 mm, height 45 mm, width 26 mm (nat. size).

Fig. 4. As above. Length 98 mm, height 58 mm, width 35 mm (nat. size). Atlantic Slope Unionidae • Johnson 417

'- *% 418 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 7

Elliptio [Elliptio) fraterna (Lea)

District River South Carolina. USNM 85396. Fig. 1. Un/o traternus Lea. Abbeville [Savannah drainage], Holotype Length 60 mm, height 31 mm, width 16 mm (slightly reduced).

USNM 85398. Fig. 2. Unio Iraternus Lea. [Chattahoochee River] Columbus [Muscogee County], Georgia. Pararype Length 69 mm, height 34 mm, width 17 mm (slightly reduced).

58 35 width 22 mm Fig. 3. Unio anthonyi Lea. Florida. Holotype USNM 84986. Length mm, height mm, (slightly reduced).

186967. 75 38 Fig. 4. Flat Creek, 8 mi. SW of Samson, Geneva County, Alabama. MCZ Length mm, height mm, width 20 mm.

8 mi. of Miller Cross Holmes Fig. 5. Elliptio mcmichaeli Clench and Turner. Choctawhatchee River, W Roads, County, Florida. Holotype MCZ 191922. Length 91 mm, height 50 mm, width 27 mm.

Elliptio waccamawens/s (Lea)

84437. 36 Fig. 6. Unio waccamawens/s Lea. Lake Waccamaw, North Carolina. Holotype USNM Length mm, height 17 mm, width 16 mm (nat. size).

North Carolina. MCZ Fig. 7. Canal beside Lake Waccamaw, 1 mi. NNW of Dupree Landing, Columbus County, 214235. Length 89 mm, height 45 mm, width 28 mm (approximately nat. size). Atlantic Slope Unionidae • Johnson 419 420 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 8

Elliptio (Elliptio) complanata (Lightfoot)

269257. 99 50 Fig. 1. Savannah River, 7 mi. NE of Newington, Screven County, Georgia. MCZ Length mm, height

mm, width 28 mm (slightly enlarged).

Wilkinson MCZ 234355. Fig. 2. Turkey Creek, 4 mi. NE of Allentown [Altamaha River system], County, Georgia. Length 73 mm, height 36 mm, width 18 mm (nat. size).

North Carolina. USNM Fig. 3. Unio quadrilaterus Lea. Neuse River, near [6 mi. E of] Raleigh, [Wake County], Holotype 85385. Length 60 mm, height 36 mm, width 20 mm (slightly reduced).

North Holo- Fig. 4. Unio roanokensis Lea. Roanoke River [road between Norfolk, Virginia, and Tarborough, Carolina]. type USNM 85423. Length 119 mm, height 57 mm, width 29 mm (slightly reduced).

North Carolina. USNM 85611. Fig. 5. Unio mediocris Lea. Neuse River, [6 mi. E of] Raleigh [Wake County], Holotype

Length 57 mm, height 30 mm, width 19 mm (slightly reduced).

North Carolina. 258813. Fig. 6. North shore, Lake Waccamaw, town of Lake Waccamaw, Columbus County, MCZ Length 60 mm, height 31 mm, width 18 mm (approximately nat. size). Atlantic Slope Unionidae • Johnson 421 422 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 9

Elliptio (Elliptic*) hopetonensis (Lea)

Macon USNM 85531. Length 57 Fig. 1. Unio inusitatis Lea. Swift Creek, below [Bibb County], Georgia. Holotype mm, height 32 mm, width 19 mm (slightly reduced).

near Darien USNM Fig. 2. Unio hopetonensis Lea. [Altamaha River], Hopeton, [Mcintosh County], Georgia. Holotype 85391. Length 84 mm, height 43 mm, width 20 mm (slightly reduced).

Elliptio [Elliptio] icjerina (Conrad)

ANSP 41381. Fig. 3. Unio icterinus Conrad. Savannah River, opposite Augusta [Richmond County], Georgia. Holotype Length 55 mm, height 28 mm, width 18 mm (approximately nat. size).

Darien USNM Fig. 4. Unio lugubris Lea. [Altamaha River], Hopeton, near [Mcintosh County], Georgia. Holotype 85638. Length 53 mm, height 37 mm, width 22 mm (slightly enlarged).

South Carolina. USNM 85669. Fig. 5. Unio fuomeyi Lea. Abbeville District [Savannah River drainage]. Holotype Length 65 mm, height 33 mm, width 19 mm (slightly reduced).

85546. 66 37 Fig. 6. Unio opacus Lea. Buckhead Creek, Burke County, Georgia. Holotype USNM Length mm, height mm, width 24 mm (slightly reduced).

South Carolina. USNM Fig. 7. Unio hastatus Lea. New Market, Abbeville District [Savannah River drainage], Holotype 86013. Length 66 mm, height 28 mm, width 17 mm (slightly reduced).

85646. 65 Fig. 8. Unio obnubilus Lea. Buckhead Creek, Burke County, Georgia. Holotype USNM Length mm, height

38 mm, width 23 mm (slightly reduced).

85653. 57 31 Fig. 9. Unio similis Lea. Buckhead Creek, Burke County, Georgia. Holotype USNM Length mm, height

mm, width 20 mm (slightly reduced).

85561. 66 Fig. 10. Unio aequotus Lea. Buckhead Creek, Burke County, Georgia. Holotype USNM Length mm, height

38 mm, width 21 mm (slightly reduced). Atlantic Slope Unionidae • Johnson 423 424 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 10

EUiptio [Elliptio] icterina (Conrad)

85077. 44 Fig. 1. Un/'o micans Lea. Catawba River, Gaston County, North Carolina. Holotype USNM Length mm, height 23 mm, width 14 mm (slightly reduced).

85901. Fig. 2. Un/'o ocmu/geens/'s Lea. Little Ocmulgee River, Lumber City, [Telfair County], Georgia. Holotype USNM Length 104 mm, height 50 mm, width 35 mm (slightly reduced).

85634. 60 Fig. 3. Un/'o confertus Lea. Santee Canal, South Carolina, from J. Lewis. Metatype USNM Length mm, height 33 mm, width 22 mm (slightly reduced).

The holotype was not located. This specimen, identified by Lea, is almost the same width and height as the original specimen, though the latter was more inflated, measuring 28 mm in width.

Elliptio [EUiptio] arctata (Conrad)

width 14 Fig. 4. Un/'o arcfatus Conrad. Alabama River, Alabama. Lectotype ANSP 41356. Length 55 mm, height 22 mm, mm (nat. size).

Fig. 5. Un/'o sfrigosus Lea. Chattahoochee River, Columbus [Muscogee County], Georgia. Holotype USNM 85890.

Length 56 mm, width 24 mm, height 13 mm (slightly reduced).

86155. Fig. 6. Un/'o lazarus Lea. Abbeville District [Savannah River drainage], South Carolina. Holotype USNM

Length 52 mm, height 23 mm, width 12 mm (slightly reduced).

Fig. 7. Un/'o perstriatus Lea. Abbeville District [Savannah River drainage], South Carolina. Holotype USNM 85892.

Length 57 mm, height 25 mm, width 12 mm (slightly reduced).

71 Fig. 8. Un/'o graci/entus Lea. Catawba River, Gaston County, North Carolina. Holotype USNM 85976. Length mm, height 30 mm, width 13 mm (slightly reduced).

Fig. 9. Un/'o perlatus Lea. Cape Fear River, Black Rock Landing [= 2 mi. S of Kings Bluff, Bladen County], North Caro- lina. Holotype USNM 86006. Length 66 mm, height 22 mm, width 12 mm (slightly reduced).

EUiptio [EUiptio] lanceolata (Lea)

Fig. 10. Un/'o productus Conrad. Savannah River, Augusta [Richmond County, Georgia]. Holotype ANSP 41397. Length 66 mm, height 26 mm, width 16.5 mm (approximately nat. size). Atlantic Slope Unionidae • Johnson 425 426 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE n

Elliptio [Elliptio] lanceolata (Lea)

85863. Fig. 1. Unio subcylindraceus Lea. Rocky Creek, near Macon [Bibb County], Georgia. Holotype USNM Length

95 mm, height 41 mm, width 36 mm (slightly reduced).

Fig. 2. Unio lanceolatus Lea. Tar River, Tarborough [Edgecombe County], North Carolina. Holotype USNM 85905.

Length 41 mm, height 18 mm, width 11 mm (slightly reduced).

29 Fig. 3. Unio angustatus Lea. Cooper River, South Carolina. Holotype USNM 85896. Length 72 mm, height mm, width 17 mm (slightly reduced).

Fig. 4. Un/o emmonsii Lea. Roanoke River, Weldon [Halifax County], North Carolina. Holotype USNM 86028.

Length 110 mm, height 46 mm, width 26 mm (slightly reduced).

Fig. 5. Unio hazelhurstianus Lea. Satil I a River, Camden County, Georgia. Holotype USNM 86009. Length 78 mm, height 35 mm, width 22 mm (slightly reduced).

Fig. 6. Unio fisherianus Lea. Head of Chester River [Kent County], Maryland. Holotype USNM 86002. Length 70 mm, height 29 mm, width 15 mm (slightly reduced).

Elliptio [Elliptio] shepardiana (Lea)

234042. 127 29 Fig. 7. Ocmulgee River, 1 mi. S of Lumber City, Telfair County, Georgia. MCZ Length mm, height mm, width 15 mm (slightly enlarged). Atlantic Slope Unionidae • Johnson 427 428 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 12

Uniomerus tetralasmus (Say)

85366. 82 Fig. 1. Unio obesus Lea. Little Ogeechee River [Hancock County], Georgia. Holotype USNM Length mm, height 50 mm, width 35 mm (slightly reduced).

85373. Fig. 2. Unio bisse/zanus Lea. Bissels Pond, Charlotte [Mecklenburg County], North Carolina. Holotype USNM Length 69 mm, height 40 mm, width 24 mm (slightly reduced).

85376. Fig. 3. Unio squalidus Lea. Roanoke River, near Weldon [Halifax County], North Carolina. Holotype USNM Length 52 mm, height 30 mm, width 18 mm (slightly reduced).

41 25 width Fig. 4. Unio cicur Lea. Little Ocmulgee River, Georgia. Holotype USNM 85532. Length mm, height mm,

17 mm (slightly reduced).

Fig. 5. Unio ineptus Lea. Abbeville District [Savannah River drainage], South Carolina. Holotype USNM 85326. Length 33 mm, height 18 mm, width 10 mm (slightly reduced).

Carolina. Fig. 6. Unio pawensis Lea. Beaver Creek [into ?], Catawba Run [Gaston and Mecklenburg Counties], North Holotype USNM 85380. Length 74 mm, height 42 mm, width 29 mm (slightly reduced).

Lasmigona (P/afyna/as) subviridis (Conrad)

Fig. 7. Unio pertenuis Lea. Neuse River, near [6 mi. E of] Raleigh [Wake County], North Carolina. Holotype USNM 86139. Length 35 mm, height 22 mm, width 13 mm (slightly reduced).

Fig. 8. Unio hyalinus Lea. [James River drainage], Richmond [Henrico County], Virginia. Holotype USNM 86131

Length 35 mm, height 21 mm, width 12 mm (slightly reduced).

Fig. 9. Unio charlottensis Lea. [Sugar Creek], near Charlotte, Mecklenburg County, North Carolina. Holotype USNM

85402. Length 101 mm, height 58 mm, width 34 mm (slightly reduced). Atlantic Slope Unionidae • Johnson 429 430 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 13

Lasmigona (Platynaias) subviridis (Conrad)

District River South Carolina. USNM 83972. Fig. 1. Un/o decorafus Lea. Abbeville [Savannah drainage], Holotype Length 60 mm, height 37 mm, width 19 mm (slightly reduced).

Alasmidonta [Prolasmidonta] heterodon (Lea)

231191. 34 19 Fig. 2. Neuse River [6 mi. E of] Raleigh [Wake County], North Carolina. MCZ Length mm, height mm, width 14 mm (1.5 X).

Alasmidonta {Alasmidonta) undulata (Say)

237455. 54 32 Fig. 3. Nottoway River, 3 mi. E of Rowlings, Brunswick County, Virginia. MCZ Length mm, height mm, width 23 mm (approximately nat. size).

29 19 Fig. 4. James River, opposite Maidens, Goochland County, Virginia. MCZ 261313. Length mm, height mm, width 13 mm (1.5 X)-

Alasmidonta (Alasmidonta) triangulata (Lea)

86249. 57 Fig. 5. Margan'fana triangulata Lea. Upper Chattahoochee [River], Georgia. Holotype USNM Length mm, height 38 mm, width 31 mm (nat. size).

48 32 width 27 Fig. 6. Mill Race, 2 mi. N of Sardis, Burke County, Georgia. MCZ 237453. Length mm, height mm, mm (slightly enlarged).

Alasmidonta (Alasmidonta) arcu/a (Lea)

86170. Fig. 7. Margaritana areola Lea. Altamaha [River], Liberty [now Long] County, Georgia. Holotype USNM Length 57 mm, height 44 mm, width 41 mm (nat. size).

50 43 Fig. 8. Altamaha River, 4 mi. NE of Jesup, Wayne County, Georgia. MCZ 237551. Length mm, height mm, width 38 mm (slightly enlarged). Atlantic Slope Unionidae • Johnson 431 432 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 14

Alasmidonta (Decurambis) varicosa (Lamarck)

44 Fig. 1. Turkey Creek, 8 mi. NW of Edgefield, Edgefield County, South Carolina. MZUM 58024. Length mm, height

26 mm, width 18 mm (slightly reduced).

216721. 45 Fig. 2. North Fork of Shenandoah River, E of Woodstock, Shenandoah County, Virginia. MCZ Length mm, height 25 mm, width 18 mm (2X)-

Anodonfa (Pyganodon) cataracta cataracta Say

from the Fig. 3. Anodonta doliaris Lea. Stewarts Mill Dam, Union County, North Carolina. Topotype MZUM 103859, 111 67 original lot collected by C. M. Wheatley. Lea saw only the holotype in the ANSP 126522a. Length mm, height mm, width 50 mm (slightly reduced).

86600. Fig. 4. Anodonfa dariensis Lea. Swift Creek, near Macon [Bibb County], Georgia. Holotype USNM Length

107 mm, height 58 mm, width 49 mm (slightly reduced). Atlantic Slope Uxionidae • Johnson 433 434 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 15

Anodonta [Pyganodon] cataracta cataracta Say

Fig. 1. Anodonta virgulata Lea. Roanoke River, Weldon [Halifax County], North Carolina. Holotype USNM 86593. Length 78 mm, height 44 mm, width 33 mm.

Anodonta [Pyganodon] gibbosa Say

Fig. 2. Altamaha River, "Riverside Park," 4 mi. N of Jesup, Wayne County, Georgia. MCZ 234056. Length 40 mm, 3 height 30 mm, width 22 mm (l /s X)-

Fig. 3. Cedar Creek, Fountains Mill, 7 mi. SW of Hawkinsville, Pulaski County, Georgia. MCZ 111438. Length 107 mm, height 61 mm, width 48 mm (slightly reduced).

Anodonta (Pyganodon) implicata Say

Fig. 4. Anodonta newtonensis Lea. Schuylkill [River], Fairmount [Philadelphia, Philadelphia County, Pennsylvania].

Lectotype USNM 86561. Length 97 mm, height 52 mm, width 39 mm (slightly reduced). Atlantic Slope Unionidae • Johnson 435 436 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 16

Anodonta (Pyganodon) implicata Say

' ' near Fairfield Con- Fig. 1. Anodonta housalonica Linsley Gould. Housatonic [River], Corum [= Huntington, County], necticut. Holotype USNM 678302. Length 94 mm, height 51 mm, width 34 mm (slightly reduced).

Massachusetts. Fig. 2. Anodonta implicata Say. Agawam River (outlet of Halfway Pond), Plymouth [Plymouth County], Neotype MCZ 176769. Length 120 mm, height 60 mm, width 48 mm (somewhat reduced). Interior view, showing the especially distinct pallial line caused by the characteristic anterior-ventral thickening of the shell.

Anodonta (Utterbackia) imbecilis Say

42 width 34 Fig. 3. Magnolia Springs, Perkins, Jenkins County, Georgia. MCZ 234050. Length 90 mm, height mm, mm (nat. size).

Anodonta [Utterbackia) couperiana Lea

50 width 35 mm Fig. 4. Pond near Savannah, Chatham County, Georgia. MCZ 119137. Length 87 mm, height mm,

(nat. size). Atlantic Slope Unionidae • Johnson 437 438 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 17

Anodonta [Utterbackia] couperiana Lea

Fig. 1. Anodonta dunlapiana Lea. [Charleston, Chatham County], South Carolina. Lectotype USNM 86564. Length 83 mm, height 46 mm, width 35 mm (slightly reduced).

Sfroph/fus undulatus (Say)

Fig. 2. Turkey Creek, 8 mi. NW of Edgefield, Edgefield County, South Carolina. MCZ 82807. Length 47 mm, height 29 mm, width 17 mm (nat. size).

Fig. 3. As above. Length 74 mm, height 44 mm, width 27 mm (nat. size).

Carunculina pulla (Conrad)

Fig. 4. Unio pullus Conrad. Wateree River, South Carolina. Type lost. (Figures after Conrad). Length 33 mm, height 19 mm, width 13.5 mm (nat. size).

Fig. 5. Caruncu/ina patrickae Bates. Savannah River, Johnsons Landing, 10 mi. W of Allendale, Allendale County South Carolina. Topotype MCZ 255220. Length 32 mm, height 19 mm, width 15.5 mm. Female (1.25 X)- This topotype lacks the sharp posterior ridge of the holotype.

Fig. 6. As above. Length 22 mm, height 13.5 mm, width 10 mm. Male (1.25 X)- This topotype closely resembles Bates' sketch of the allotype.

Fig. 7. University Lake, an impoundment on Morgan Creek, 1 mi. W of Chapel Hill, Orange County, North Carolina. MCZ 261347. Length 25 mm, height 17 mm, width 11 mm. Male (1.25 X).

Villosa villosa (Wright)

Fig. 8. St. Marys River, Nassau County, Florida. USNM 152066. Length 56 mm, height 29 mm, width 20 mm. Male

(slightly reduced).

Fig. 9. Unio villosus Wright. Suwannee River [Luraville], Suwannee County, Florida. Lectotype USNM 150503. Length

57 mm, height 28 mm, width 18 mm. Female (slightly reduced).

V///osa vibex (Conrad)

Fig. 10. Unio vibex Conrad. Black Warrior River, S of Blount s Spring [Blount County], Alabama. Holotype ANSP 56488a. Length 49 mm, height 28 mm, width 17 mm. Male (approximately nat. size).

Fig. 11. As above. Allotype ANSP 56488. Length 54 mm, height 31 mm, width 19 mm. Female (approximately nat. size). Atlantic Slope Unionidae • Johnson 439

- / 10 440 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 18

Villosa vibex (Conrad)

Fig. 1. Um'o gracilior Lea. Buckhead Creek [Burke County]; or Tobesaufke [Tobesofkee] Creek, near Macon [Bibb

County],- both Georgia. Holotype USNM 85088 [exact locality not indicated]. Length 50 mm, height 28 mm, width 18 mm. Male (slightly reduced).

Fig. 2. Um'o modioliformis Lea. Santee Canal, South Carolina. Probable holotype USNM 85029. Length 68 mm, height 40 mm, width 25 mm. Female (slightly reduced).

Fig. 3. Mill Race, 2 mi. N of Sardis, Burke County, Georgia. MCZ 234263. Length 48 mm, height 27 mm, width 17 mm. Male (nat. size).

Villosa delumbis (Conrad)

Fig. 4. Un/'o concavus Lea. Abbeville District [Savannah River drainage], South Carolina. Holotype USNM 85154.

Length 55 mm, height 32 mm, width 21 mm. Male (slightly reduced).

Fig. 5. Um'o genui'nus Lea. Bissels Pond, Charlotte [Mecklenburg County], North Carolina. Holotype USNM 85123.

Length 52 mm, height 30 mm, width 20 mm. Male (slightly reduced).

Fig. 6. Mill Race, 2 mi. N of Sardis, Burke County, Georgia. MCZ 234340. Length 53 mm, height 30 mm, width 19 mm. Male (nat. size).

Fig. 7. As above. Length 57 mm, height 36 mm, width 19 mm. Female (nat. size).

Fig. 8. Um'o ogeecheensi's Conrad. Ogeechee River, Georgia. Lectotype MCZ 146971. Length 64 mm, height 39 mm, width 30 mm. Female (approximately nat. size).

Villosa constricta (Conrad)

Fig. 9. Um'o lienosus constrictus Conrad. North [:= Maury] River, Rockbridge County, Virginia. Lectotype ANSP 56465a. Length 41.4 mm, height 27 mm, width 16 mm. Female (approximately nat. size).

Fig. 10. Um'o genthii Lea. [Probably from] Deep River, Gulf [Chatham County], North Carolina. Holotype USNM 84834. Length 39 mm, height 27 mm, width 18 mm. Male (nat. size). Atlantic Slope Uxionidae • Johnson 441 442 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 19

Ligumia nasuta (Say)

Fig. 1. Potomac River, Washington, D.C. MCZ 119087. Length 62 mm, height 25 mm, width 12 mm. Male.

Fig. 2. As above. MCZ 5653. Length 66 mm, height 29 mm, width 14 mm. Female.

Lampsilis [Lampsilis) can'osa (Say)

Fig. 3. Savannah River, 7 mi. NE of Newington, Screven County, Georgia. MCZ 234241. Length 98 mm, height 62 mm, width 42 mm. Male (slightly reduced).

Fig. 4. As above. Length 98 mm, height 69 mm, width 45 mm. Female (slightly reduced).

Fig. 5. Un/'o crocatus Lea. Savannah River, Georgia. Holotype USNM 84908. Length 42 mm, height 27 mm, width 19 mm. Male (slightly reduced). Atlantic Slope Unionidae • Johnson 443 444 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 20

Lampsilis [Lampsilis) dolabraelormh (Lea)

Fig. 1. Altamaha River, 11 mi. N of Odum, Wayne County, Georgia. MCZ 234197. Length 16 mm, height 12 mm, width 7.5 mm. Male (4X1-

Fig. 2. Altamaha River, 10 mi. N of Baxley, Appling County, Georgia. MCZ 234010. Length 91 mm, height 69 mm, width 49 mm. Female (slightly reduced).

Fig. 3. As above. Length 79 mm, height 63 mm, width 50 mm. Male (slightly reduced).

Fig. 4. Unio dolabraeformh Lea. Altamaha River, Liberty [now Long] County, Georgia. Holotype USNM 84888. Length 116 mm, height 79 mm, width 51 mm. Male (reduced). Atlantic Slope Unioxidae • Johnson 445 446 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 21

Lampsilis- [Lampsilis] ovata (Say)

Lecto- Fig. 1. Lampsilis ventricosa cohongoronfa Ortmann. Potomac River, Hancock, Washington County, Maryland. type Carnegie Museum 61.3999. Length 95 mm, height 60 mm, width 41 mm. Male.

width 42 mm. Female. Fig. 2. As above. Allotype Carnegie Museum 61.4000. Length 87 mm, height 62 mm,

Lampsilis [Lampsilis] ochracea (Say)

44 width Fig. 3. Un/'o rosaceus Conrad. Savannah River, Georgia. Holotype MCZ 178779. Length 73 mm, height mm, 27 mm. Male (approximately nat. size).

40 Fig. 4. Lake Waccamaw, [town of] Lake Waccamaw, Columbus County, North Carolina. MCZ 234327. Length mm, height 30 mm, width 17 mm. Female (approximately nat. size).

Fig. 5. As above. Length 50 mm, height 35 mm, width 21 mm. Male (approximately nat. size). Atlantic Slope Unionidae • Johnson 447 448 Bulletin Museum of Comparative Zoology, Vol. 140, No. 6

PLATE 22

Lampsilis (Lampsilis) radiata radiata (Gmelin)

Fig. 1. Unio conspicuus Lea. Yadkin River, Salisbury [Rowan County], North Carolina. Holotype USNM 85056. Length 112 mm, height 64 mm, width 39 mm. Male (slightly reduced).

523976. 70 Fig. 2. Greenfield Pond, Wilmington [New Hanover County], North Carolina. USNM Length mm, height

43 mm, width 28 mm. Male (slightly reduced).

width 28 Female Fig. 3. As above. USNM 452041. Length 76 mm, height 40 mm, mm. (slightly reduced).

Lampsilis (Lampsilis) splendida (Lea)

234019. 71 47 Fig. 4. Ocmulgee River, 1 mi. S of Lumber City, Telfair County, Georgia. MCZ Length mm, height mm, width 35 mm. Female (approximately 1.2 X).

84893. Fig. 5. Unio splendidus Lea. Alfamaha River [Mcintosh or Long County], Georgia. Holotype USNM Length 70 mm, height 43 mm, width 36 mm. Male (slightly reduced). Atlantic Slope Unionidae • Johnson 449