bobda bind Mates tila tated Raa es 1t sibrhy!) fails ‘ Prue cul (i tat Ahi deity Wi Me bilgi ear teotgtated viaits's tai } Sha piplii phil \ Hf yet babel gay a eta Latellels arian peasant it 4 i Aedes BUH ES , f i nH Lt Avid f RAPE EL! oe Tethalt i Ve bails PURE tenet it A shidetaitay APS CRYIN Petale balks Pala haaad at ita bele Vp Let eats PR TH tytek 7 ! het Lib f} rf git if. ain elheitallal A NT ne tie vita Ndatehe 1 by “4 lt Hy \ args? anita t} ifiial pian H it a ar Beta Nt ae i as hops ait Ta liditananay hy ie an wee pe rth] DEAT UE RN 1 5 4 i ' uh THI I if aay ty f laa sialsetey if yh prlete POruneent gatateel Fails p sdanitanited reyes ini iy bedat bt hy teat itt state , TREE NEY a i oN telah i i ic Aah ee ith He Hr fe Niet t ea es < titel 8 « a thee iN Hit ch a rane ch H iat + uate i nied ie o as iit us tat SH be tate Ware tats! aie Ponies cas if Tale ta ye Venues dated HN inf i) HM i A He iat uly Hat ily Me | | iM tt ah i Tait bh ae iit t dun ded nhs Naaaalet H Aida dl Gs Te eines Nettie t ben} sadaine tet int i 14 Ae HBL Ue ‘ fhe $ t H heaaaay'y ay Parere rns ieee ty ai EV Alniy nie 7 +1 Date ts AM) eat oh it it 7 teen aNd Wish oy ite ‘ ehitwnad ay He inh ss att ae ie i ATO Lae y dy fj ie u] aot ite Hay OED Sc sae ieiahssaansiany repens Hight Ries utlabinnae steaks Hane An! uit He “4 te H Cie Ha i ate fh aa | ai he an ae tet wae Hy Pit Ai \] Heh hit M mes ; isi rans EX LIBRIS William Healey Dall Division of Mollusks Sectional Library A ea oe Ra ey “ Sonderabdruck »Aus der Natur“, Jahrg. 1907/8. OL. Geeiondl Library Nordamerikanische FluBmuscheln. Von Dr. A.E.ORTMANN, Curator of Zoology, Carnegie Museum, Pittsburgh, Pa. (Mit zw6lf Abbildungen.) Eines der am stirksten und ausgedehntesten entwickelten Flufisysteme der Welt ist das des Mississippi in Nordamerika, und dementsprechend haben auch viele Tiergruppen, die dem Siifwasser angehéren, in Nordamerika eine ungemeine Mannigfaltigkeit und Reichhaltigkeit erreicht. Dies gilt z. B. fiir die Schildkréten, die geschwinzten Amphibien, fiir Fische; aber auch manche Wirbellosen, wie z. B. die Flu{krebse und besonders die Mollusken. Unter letzteren sind es besonders zwei Familien, die hier einen Reichtum der Entwicklung aufweisen, wie er auf der ganzen tibrigen Welt beispiel- los ist: es sind dies die den Gastropoden oder Schnecken angehorige Familie der Strepomatidae (neuweltlichen Melanien), und die eigentlichen FluBmuscheln, die Unionidae. Von letzteren sind etwa tausend Arten aus allen Kontinenten bekannt; davon besitzt allein Nordamerika tber finf- hundert; der bei weitem gréfte Teil findet sich im Mississippi-System, nur verhiltnismibig wenige in den Fliissen, die dem Atlantischen Ozean (6st- lich von der Mississippimiindung, zustr6men und noch weniger auf der pazifischen Seite der Felsengebirge. Der Verfasser hatte in den letzten Jahren Gelegenheit mit der Fluf- muschelfauna des Mississippigebietes naher bekannt zu werden, und zwar im oberen Ohiobecken, im westlichen Pennsylvanien. Wenn auch in dieser Gegend die Fauna schon etwas verarmt ist, so war er doch imstande, hier nicht weniger wie 45 Arten nachzuweisen, eine Zahl, die sicherlich durch weitere Forschungen noch vermehrt werden wird. Es ist durchaus nicht auBergewohnlich, auf einer einzigen Exkursion an giinstigen Stellen der groferen Fliisse (Ohio, Allegheny) etwa 20 Arten in engster Nachbarschaft zu finden, ein Verhialtnis, das ohne weiteres die Reichaltigkeit der Fauna bezeugt, wenn wir z. B. damit die geringe Zahl der Arten vergleichen, die sich in ganz Deutschland auffinden lassen. Den allgemeinen Bau der FluSmuscheln diirfen wir wohl als bekannt voraussetzen. Hier soll nur ein Punkt hervorgehoben werden, der syste- matisch auferordentlich wichtig ist, aber erst in verhaltnismafig neuer Zeit in seiner Bedeutung erkannt: wurde (vgl. diese Zeitschr. Jg. IL, S. 891). Seitlich vom Fu der Muschel, zwischen diesem ung a en} Fed ae fos 2 Dr. A. E. ORTMANN: lappen, die der Innenseite der Schale anliegen und diese abscheiden, finden sich jederseits zwei lamellenartige Organe, die Kiemen. Sie dienen in erster Linie zur Atmung. Nun tritt aber hier zu gewissen Zeiten ein eigen- tiimlicher Funktionswechsel auf. bei der weiblichen!) Muschel wandern -. gur Vortpilanzungszeit die Eier in die Kiemen hinein und beginnen dort ihre Entwicklung, bis sie, nach Erreichung eines Embryonalstadiums, Glochi- diyn genannt, ins Wasser entleert werden. Die mit Eiern oder Embryonen gefiiiltle Kieme, oder Teile derselben, wird Marsupiuwm genannt, und die Marsupien werden zur Einteilung der FluBmuscheln in erster Linie herangezogen. Friher stiitzte sich das System der Flu{muscheln vorwiegend, ja aus- -schlieBlich auf die Schalen. Die ungeheure Formenmannigfaltigkeit der letzteren gab zu verschiedenen Einteilungen Veranlassung, jedoch befriedigte keime derselben, und man kam immer wieder auf die alte Einteilung in zwei oder drei Gattungen zuriick. Man nannte solche Formen, die gut ent- wickelte Schlofizihne besitzen, Unio, und solche, bei denen letztere fehlen, Anodonta. Dazwischen stellte man dann gewéhnlich eine dritte Gattung mit mehr oder weniger reduzierten Schlofizihnen (Margaritana oder Alasmi- donta). Abgesehen davon, dai diese Gattungen, besonders Unzo, einen un- geheuren Umfang hatten, entsprach diese Einteilung nicht den natiirlichen Verwandtschaftsverhaltnissen: wir wissen jetzt, dafi der Verlust der SchloB- zihbne unabhingig voneinander in verschiedenen Gruppen stattfand, und dafi es deshalb nicht angiingig ist, die mit reduzierten Schlofzaihnen ver- sehenen Formen in nur einer oder zwei Gattungen unterzubringen. Mit Beriicksichtigung der Weichteile und besonders des Marsupium des Weibchens ist es jedoch mdglich, eine naturgemafe Einteilung an — Stelle der alten, unnatiirlichen zu setzen. Bahnbrechend in dieser Richtung war eine Arbeit von C. T. Simpson?), dem es gelang, ein System auszuarbeiten, das, wenn auch noch nicht in allen Einzelheiten einwandfrei, doch die Hauptgesichtspunkte andeutet, nach denen man zu einem Verstandnis der ver- wandtschaftlichen Beziehungen gelangen kann. Das Hauptkriterium hierbei wird aber vom Marsupium geliefert®). Wir kénnen auf die Einzelheiten hier nicht naher eingehen. Nur so viel soll gesagt sein, dafi ein allgemeiner Fortschritt in der Entwicklung des Masurpiums deutlich zu erkennen ist. bei den primitivsten Formen (Gattung Quadrula) werden alle vier Kiemen zu Eibehaltern. Bei allen iibrigen Gattungen ist es nur das duferste Paar, welches zunichst ganz als Marsupium dient (z. B. Unio, Margaritana, Alasmidonta, Symphynota, Anodonta u. a.). Von diesem Punkt an beginnen ') Bei den meisten FluBmuscheln sind die Geschlechter getrennt, doch finden sich auch hermaphroditische Formen. Von einer Art (Anodonta imbecillis Say) ist letzteres mit Bestimmtheit (durch Dr. V. SrerKi) nachgewiesen worden. *) C. T. Simpson, Synopsis of the Najades, or pearly freshwater Mussels, in: Proceed. U. S. Mus. 22. 1900. ®) Es mufi ganz entschieden hervorgehoben werden, dai Simpsons System keines- wegs allein auf das Marsupium gegriindet ist. Andere Charaktere, der Weichteile so- wohl als auch Schalencharaktere, sind eingehend beriicksichtigt worden. Doch hat sich eben herausgestellt, dafi gerade die letzteren hiufig konvergent entwickelt sind, d. h. in- einander fernstehenden Gruppen unabhingig erworben wurden. Nordamerikanische FluSmuscheln. 3 offenbar mehrere Entwicklungsrichtungen. Alle iibrigen Gattungen haben gemein, das die Kier in eine Anzahl »Kiersacke« cingeschlossen werden, die teils quer durch die Kieme verlaufen (Gattung Sfrvp/ztus), teils vertikal (dorso-ventral) oder schrig gerichtet sind, und aufierdem sind die Kiersicke auBerlich durch Furchen markiert.. Das Marsupium nimmi ‘ann ferner eine mehr und mebr lokalisierte Lage ein. Entweder liegt es am ventralen Rand der Kieme (Dromas und Ptychobranchus, bei letzterer ist es cicentiimlich gefialtelt); oder es liegt in der Mitte der Kieme (Obliquaria, Cy; ogenia), oder im hinteren Abschnitt derselben (eine ganze Reihe Gattungen, die wichtigsten: sind Lampsilis und Truncilla). Diese letztere Gruppe ist offen- bar die am héchsten entwickelte; dies geht auch aus einem eigentiimlichen Verhiltnis in den. Schalen hervor. Wéahrend bei allen itbrigen Formen minnliche und weibliche Tiere (wenn die Geschlechter verschieden sind) sich 4uferlich nicht unterscheiden lassen, tritt hier eine sexuelle Differenzie- rung der Schalen ein. In der Gegend, wo das Marsupium liegt, findet sich - beim Weibchen eine eigentiimliche Schwellung, Verbreiterung oder Ver- Abb. la. Lampsilis luteola (Lam.) '/s. Abb. 1b. Lampsilis luteola (Lam.) */s. (Mannchen). (Weibchen). Mahoning River, Lawrence Co., Pa. Mahoning River, Lawrence Co, Pa. langerung der Schale, oft verbunden mit einer Abnahme von Dicke, -die den auferen Umrili der Schale von dem des Mannchens oft ganz ver- schieden erscheinen lat (siehe Abb. 1a und 1b, Abb. 2a bis 2d). Ehe dies Verhaltnis erkannt war, wurden hiufig Mannchen und Weibchen der betreffenden Arten verschieden beschrieben. Bei den Arten der Gattung Truncilla, wo die sexuelle Differenzierung der Schalen am ausgesprochensten ~ ist, werden aufierdem beim Weibchen am hinteren Schalenrande ineinander- greifende Zahnchen entwickelt, wie sie z.B. bei den Herzmuscheln (Cardzwm) bekannt sind, eine Erscheinung, die unter den FluBmuscheln einzig dasteht (siehe Abb. 2c und 24d). Dafi die soeben geschilderten Verinderungen des Marsupiums eine Spezialisierung in der Organisation markieren, und dali demnach ein System, das auf diese Verhiltnisse sich griindet, ein natiirliches genannt werden muh, diirfte klar sein. Hand in Hand damit gehen nun gewisse Schalen- charaktere, doch la®t sich in bezug auf diese wenig allgemeines sagen. Sehr wichtig ist die Skulptur des Wirbels, und gute Gattungs- und Art- charaktere werden durch sie geliefert. Doch ist dieselbe nicht immer zu beobachten, da die Wirbel sehr haufig stark korrodiert sind; nur bei jungen Exemplaren kann man gewohnlich die Wirbelskulptur noch erhalten finden. 4 Dr. A. E. ORTMANN: Was die Schale selbst anbetrifft, so sind die primitiveren Formen (Quadrula) gewohnlich schwerer, solider, mehr oder weniger rundlich oder oval (Abb. 3), mit aufierordentlich kraftigen Schlofizihnen. Die Farbe der Epidermis ist triibe, braun oder schwiirzlich, ohne Zeichnung. Bei den héher entwickelten Formen wird im allgemeinen die Schale leichter, und es treten hiufig auf der Epidermis sehr schéne Zeichnungen auf, meist radiale Streifen, dunkel- griin auf hellgriinem oder gelblichem Grunde (siehe Abb. 1, 2, 4, 6). Oft werden die Schlofizihne reduziert, und oft nimmt die Schale eigentiimliche Gestalten an (vgl. Abb. 11, 12) und erhalt eine besondere Skulptur. Aber Abb. 2a. Truncilla triquetra (Raf.) 1/,. Abb. 2c. Truncilla triquetra (Raf.) 1/,. (Mannchen). (Weibchen). Allegheny River, Armstrong Co., Pa. Allegheny River, Armstrong Co., Pa. f Abb. 2b. Ventrale Ansicht desin Abb.2a Abb. 2d. Ventrale Ansicht des in Abb. 2c dargestellten Exemplars. dargestellten Exemplars. gerade eigentiimliche Skulpturverhaltnisse finden sich auch bei primitiven Formen (Abb. 8, 10), und sehr haufig treten primitive Formverhiltnisse noch bei héher entwickelten Arten auf (Abb. 4). Dies ist indessen durchaus nicht wunderbar. Die Schale ist der Teil der Muschel, der in ‘unmittelbare Beriihrung mit der Aufenwelt kommt, und es ist zu erwarten, dai je nach der Umgebung die Schalen ver- schiedene Anpassungen zeigen, und dafi ahnliche Anpassungen bei ver- schiedenen Formen auftreten sollten. Dies scheint nun in der Tat der Fall zu sein. Die zahlreichen Arten unseres Gebietes kommen durchaus nicht alle unter denselben Verhialtnissen vor, auch wenn sich oft eine grofie Anzahl benachbart findet. Ihr Aufenthaltsort ist der Grund der Fliisse, Str6me und Seen, wo sich die Exemplare gewohnlich ganz in ne Nordamerikanische Flu®muscheln. 5 Gerdll, Sand oder Schlamm eingraben, so dai nur das Hinterende, an dem die Ein- und Ausfuhréffnung fiir das Wasser, das zur Nahrung und Atmung dient, ein wenig herausragt. (Die Abbildungen, die diesem Auf- satz beigegeben sind, sind durchweg so gezeichnet, dai sie die natirliche Stellang der Muschel im Fluigrunde wiederzugeben suchen; in manchen Fallen sollte das Hinterteil eher noch mehr erhoben sein.) Die meisten Arten leben in flieBendem Wasser, weniger in stehendem; aber auch unter den ersteren sind die Verhiltnisse, unter denen sie sich finden, sehr mannigfach. Am haufigsten sind Muscheln an solchen Stellen der (liisse, wo sich Stromschnellen, Sandbanke und Inseln finden, wo in dei ver- schiedenen FluBarmen sehr wechselnde Bedingungen dicht nebeneinander liegen; in stirker str6mendem Wasser, mit grobem Kies als Untergrund leben hier andere Arten, als in ruhigeren Buchten mit feinerem Sand oder Schlamm. Bisher ist diesem Punkte, den dkologischen Gewohnheiten der Muscheln und den damit zusammenhingenden Gestaltungserscheinungen, Abb. 3. Quadrula subrotunda (Lea) 1/5. Abb. 4. Lampsilis ligamentina (Lam.) */s. Allegheny River, Armstrong Co., Pa. Mahoning River, Lawrence Co., Pa. wenig Aufmerksamkeit geschenkt worden. Meine eigenen Studien haben indessen, obgleich sie noch.recht unvollkommen sind, manche_interes- santen Beziehungen ergeben, und ich méchte hier auf einige derselben hinweisen. Die einfachste Gestalt einer Muschelschale ist rund oder oval im Umrif, seitlich etwas, aber nicht sehr komprimiert, ohne jede Vorspriinge und Skulptur (vgl. Abb. 3 und 4). Solche Formen, die gewohnlich recht solide und schwer sind, leben mit Vorliebe in den gr6éferen Fliissen, ein- gegraben zwischen grobem Gerdéll. Sie sind zwar nicht sehr wahlerisch und finden sich auch in ruhigerem Wasser und feinem Kies, aber sie sind jedenfalls diejenigen Arten, die grobes Ger6ll und starke StrOmung am besten ertragen. Diese Gestalt tritt in vielen Gattungen auf, und es ist klar, dafi sie eine Anpassung an die Verhiltnisse ist, unter der die Muschel lebt; sie gleicht auferlich auferordentlich einem vom Wasser gerundeten Steine und muf sich natiirlich der Str6mung gegeniiber als solcher ver- halten, d. h. von letzterer erfafit, ist es bei dieser Gestalt so gut wie ausgeschlossen, dafi die Muschel Schaden nimmt. Manche Arten sind in auffallender Weise verlingert (siehe Abb. 5, 6 u. 7). Auch sie gehéren sehr verschiedenen Gattungen an und finden : i pe 7 to, ~ a Re tie Raa oes PS SEES TEE TT EE = ; bay? LGR (a= ' man bedenkt, dafi die betreffenden Arten 2 ; 4 Pa ee » 7a 6 ' Dr. A. E. ORTMANN: eats sich oft in starker Strémung und in grobem Gerdll. Diese verlingerte . Gestalt dient walvscheinlich zur besseren Verankerung. Es ist lei at begreiflich, daf von zwei Muscheln von gleichem Vo- lumen diejenige besser befestigt ist, die langer ist, und deshalb tiefer im Kies stecki. Wihrend also die im -voranstehenden Abschnitt genannten Arten (Abb. 8 u. 4) keinen so grofen Wert auf die Verankerung legen, aber gegen die Folgen des Ergriffenwerdens von der Strémung geschitzt sind, legen diese Formen das Haupt- saat hae gewicht auf feste Verankerung, um aS ein Ergriffenwerden von. der Strémung moglichst zu vermeiden. Derselbe Zweck, Schutz gegen Zertriimmerung von der Strémung, wird also auf zwei Wegen erreicht. Eine feste Verankerung wird ferner 2 noch auf andere Weise erreicht, nim- Abb. 5. Unio gibbosus Barn. */s. lich durch Entwicklung einer Ober- Allegheny River, Armstrong Co., Pa. flachenskulptur (siehe Abb. 8, 9 u. 10). Es koénnte vielleicht daran~ gedacht werden, dafi diese Skulptur, wo sie aus Knoten, Héckern u. dgl. besteht, dazu dienen mdodchte, der Schale erhdhte Festigkeit zu verleihen; dies erscheint indessen nicht als wahrscheinlich, wenn gewohnlich schon so wie so feste, solide Scehalen besitzen, und bei diinnschaligen Abb. 6. Ptychobranchus phaseolus (Hildr.) Arten solche Skulpturen kaum bekannt ‘/2- Mahoning River, Lawrence Co., Pa. sind. Ich glaube vielmehr, dafi es sich Ses Eee lediglich um eine Vermehrung der ee ay Oberfliiche handelt, um die Reibung Drs, mit dem umgebenden Sand und r(( es Kies zu vergréfern und somit eine #(* festere Verankerung herbeizufiih- We = ren. Bei dem merkwiirdigen Unio Semen = pine Lee) ans dem Alameda Ape amps ren (a) R y Allegheny River, Armstrong Co., Pa. seite der Schale lange Dornen ent- wickelt (Abb. 9), ist diese Idee jedenfalls sehr einleuchtend. Kin eigentiimlicher Fall liegt bei Quadrula wndulata (Barn.) vor (Abb. 10). . Hier verlaufen auf der Schalenfliche mehrere grobe Wiilste, und zwar in eigentiimlicher Richtung. (Auch andere Arten zeigen ahnliches.) Wahrend es nun bei Muscheln fast eine allgemeine Regel ist, dafi etwa vorhandene _ Schalenskulptur entweder konzentrisch oder radial verliuft, nehmen hier diese Wiilste auf der breiten Schalenfliche eine intermediare, schrige Richtung ein, und nur nahe dem oberen Hinterrande werden sie nahezu Nordamerikanische Flufimuscheln. 7 radial. Bringt man aber die Muschel in die natiirliche Stellung, wie sie im Kiese steckt (so wie es in Abb. 10 geschehen ist), so sieht man, dah nun die Wiilste horizontal verlaufen, gleichlaufend init der Bewegungs- richtung der Muschel. Beim Vorwartskriechen der letzteren (und diese Art ist tatsichlich auBergewohnlich wanderlustig) bieten also die Wiilste keinen Widerstand dar, und wirken nur als Verankerungsapparat, wenn irgendeine Kraft die Muschel nach oben, aus dem Kiese heraus, zu ziehen Abb. 8» Quadrula metranevra (Raf.) 1/s. Abb. 9. Unio spinosus (Lea.) */s. Allegheny River, Armstrong Co., Pa. Altamaha River, Ga. versuchen sollte. Beim Kriechen des Tieres funktionieren diese Wiilste gewissermafien als Gleitschienen, oder etwa wie die Kufen eines Schlittens, nur daf} sie nicht unten, sondern seitlich angebracht sind. Die Kriimmung der Wiilste nach oben im hinteren Teile der Schale kénnte man vielleicht so erklaren, daf} sie dazu dient, das Tier beim Vorwiartsbewegen tiefer in den Kies einzusenken. Eine Verditinnung der Schale tritt ebenfalls bei sehr verschiedenen Gattungen auf. Sie ist charakteristisch fiir die Gattung Anodonta, findet sich aber auch in auffallendem Mabe z. B. bei Lampsilis (Proptera) gracilis (Say). Bei derartigen Formen werden auch haufig die Schlofizihne reduziert, was vielfach wohl weiter nichts bedeutet, Z als eine fernere Erleichterung der Abb. 10. Quadrula undulata (Barn.) 1/9. Schale, und ee solche Erleichterung Mahoning River, Lawrence Co., Pa. ist notig, wenneinVersinken der Muschel im weichen Untergrunde unmoglich ge- macht werden soll. In der Tat ziehen solche diinnschaligen Arten entschieden weichen, sandigen oder schlammigen Grund vor; dies gilt im hiesigen Gebiet ganz besonders von Anodonta grandis (Say), einer typischen Schlammform, die sich vorwiegend in stehenden Gewiissern findet, seltener in ruhigen, schlammigen Buchten von Fliissen. Daf schwere Muschelschalen, wenn sie zufallig auf Schlammboden geraten, dort versinken und ersticken, ist beobachtet worden. Die eben erwahnte Reduktion der Schlofizahne findet sich auch bei anderen Formen, die ziemlich schwere Schalen haben. Ihre Bedeutung 8 Dr. A. E. ORTMANN: ist noch unklar in diesen Fallen. Indessen ist es interessant, zu beob- achten, dafi oft Hand in Hand damit eine eigentiimliche Verschmelzung beider Schalen geht. Diese findet sich am oberen Hinterrande der Muschel, wo sich dann gewodhnlich eine fligelartige Verbreiterung der Schale entwickelt (siehe Abb. 11 und 12), langs deren Oberrandes die beiden Schalen fest miteinander verschmelzen. Dies ist offenbar ein Ersatz fiir den Verlust der Schlofizahne, und auch diese Einrichtung findet sich bei Arten verschiedener, in keinem naheren Verhialtnis stehender Gaitungen, wie Symphynota, Lampsilis. Auch bei Anodonta-Arten kommt eine derartige. Tendenz vor. Zum Schlufi will ich noch auf den 6konomischen Wert der Muscheln aufmerksam machen. Die Flufimuscheln sind efibar und wurden schon von den Ureinwohnern Nordamerikas, den Indianern, als Nahrung Abb. 11. Symphynota (Pterosygna) com- Abb. 12. Lampsilis (Proptera) alata planata (Barn.) 1/,. (Say) */o. Ohio River, Wheeling, W.. Va. Monongahela River, Washington Co., Pa. benutzt. Auch jetzt noch werden sie gelegentlich gegessen, doch gestattet die Zihigkeit des muskulésen Fubes eine erfolgreiche Verwendung nur zur Bereitung von Suppen. Die Schale der Flu{muscheln, deren Innenseite aus einer mehr oder weniger dicken Perlmuttersechicht besteht, wird bei der Fabrikation von Knépfen (besonders der kleinen Perlmutterknépfe an weifier Wische) verwendet. Der Hauptsitz dieser Industrie ist in Muscatine im Staate lowa, und im Mississippigebiete werden zu diesem Zweck die Muschelschalen im Grofen gefischt. SchlieBlich finden sich in den nordamerikanischen Flufmuscheln auc Perlen, die bisweilen erheblichen Wert erreichen, wenngleich sie den marinen Perlen im Durchschnitt nachstehen. Die Entstehung der Perlen ist ahnlich wie bei der marinen Perlmuschel (Pteria): sie werden gebildet durch Ausscheidung von Perlmuttersubstanz um einen Kern herum, der von kleinen Partikeln von Sand oder Schmutz geliefert wird, die zufallig in die Muschel hineingeraten (vielleicht von einem Parasiten, wie bei Pteria?). Die Perlbildung ist also eine aufiergewohnliche oder vielleicht Nordamerikanische Flufimuscheln. g selbst krankhafte Erscheinung. Verfasser hat Perlen von einiger Grofe wiederholt gefunden, und kleinere, wertlose Perlen sind nicht gerade selten, finden sich jedoch an gewissen Lokalitaéten haufiger als an anderen. Eine der wichtigsten und bekanntesten der perlenfiihrenden Arten ist Margari- tana marguritifera (L.), die zirkumpolare Verbreitung besiizt (Europa, Nord- asien, Japan, nordliches Nordamerika). Im oberen Ohiogebiete liefern (nach des Verfassers Erfahrungen) die folgenden Arten besonders haufig Perlen: Lampsilis ventricosa (Lam.), Lampsilis ligamentina (Lam.), ‘(uadrula subrotunda (Lea), Symphynota costata (Raf.), Unio crassidens (Lam.) und andere. Das Sammeln der Muscheln fiir die Knopfindustrie und das Offnen derselben, um Perlen zu finden, wird in vielen Teilen der Vereinigten Staaten teils als Geschaft, teils als Sport betrieben. Nimmt man dazu die ausgedehnte Verunreinigung des Flufiwassers in vielen Industriegegenden dureh allerlei schadliche Stoffe — besonders auch in den Kohlengebieten des westlichen Pennsylvaniens — so ist es nicht zu verwundern, daf) diese prichtige Fauna stark im Riickgange begriffen, ja aus manchen Strémen bereits véllig verschwunden ist. Der Durchschnitts-Amerikaner besitzt ja iiberhaupt kein Verstandnis fiir die Idee der »Erhaltung von Naturdenk- milern«, und so kiimmert er sich auch, nicht darum, wenn interessante — Glieder der so reichen heimischen Fauna zugrunde gehen. Indessen muf es zum Lobe der Regierung in Washington gesagt werden, dafi sie im Sinne hat, der Zerstérung der Flufimuschelfauna zu steuern. Im _ ver- gangenen Sommer wurden Erhebungen angestellt, um zu konstatieren, in welcher Weise und bis zu welchem Grade die Flufmuschelfischerei ge- trieben wird; noch nichts ist verdffentlicht worden, so dafi der Verfasser keine Einzelheiten angeben kann.') Aber diese Erhebungen haben den Zweck, wenn méglich, die Muscheln zu erhalten, und ihrer riicksichtslosen Ausrottung durch Regierungsmafiregeln Schranken zu setzen. 1) Statistische Nachweise tiber die Muschelfischerei sind von der U. S. Commission gesammelt und publiziert worden, sind aber jetzt veraltet (etwa 10 Jahre alt). Neuere Statistiken betonen nur die Schwierigkeit, zuverlassige Daten zu erlangen. ee A PRELIMINARY. LIST: OF THE UNIONIDA® OF WESTERN PENNSYLVANIA, WITH NEW LOGAELFITES FOR -SPECIES FROM EASTERN PENNSYLVANIA Dr. A. E. ORTMANN (Issued April £2, 1909.) {Reprinted from ANNALS OF THE CARNEGIE MusEUM, VOL. V, No. 2 and 3, 1909. ] Vile A PRELIMINARY LIST OF THE, UNIONIDAY OF WESTERN PENNSYLVANIA, WITH NEW LOCALITIES FOR SPECIES FROM EASTERN PENNSYLVANIA. By Dr. A. E. ORTMANN. Several years ago, the writer, in pursuance of his studies of the fresh- water fauna of Pennsylvania, began to collect the freshwater shells. The past summer (1908) was especially favorable for collecting Unionidz, on account of the low stage of the rivers, and a good deal of work was done in this group, so that it is believed that the study of the mussels has progressed far enough to make it possible to give a general account of the Unionidz of the western half of the state. The region investigated covers not only the drainage of the upper Ohio (Monongahela and Allegheny Rivers), but also that of Lake Erie, and some of the tributaries of the Susquehanna. ‘The latter have not, however, been as thoroughly studied as the streams lying to the west of the divide. A number of additional localities are situated in southeastern Pennsylvania. Before we give our list of species, it seems to be weli to mention and discuss the previous records of the Unionidz of western Pennsylvania, since in many cases the nomenclature has changed, and since several incorrectly identified species have been listed. In the older publications we have only a few records from our region. I was only able to find the four following : Unio zsopus Green (Contr. Maclur. Lyc., I, 1827) = Pleurobema @sopus (Green). Described from the ‘‘ rivers in the neighborhood of Pittsburgh.’’ Stil present in the Ohio below, and in the Allegheny above Pittsburgh. Unio circulus Lea (Observ., I, 1834) = Obovaria circulus (Lea). Cited by Lea from the Monongahela River at Pittsburgh. The locality is now barren, no Unionidz being any longer present in the immediate vicinity of Pittsburgh. Nevertheless a few years ago this species was still present in the Ohio River below Pittsburgh. Unio coccineus Conrad (Monogr., 3, 1836, and Lea, Observ., II, 1838) = Quadrula coccinea (Conrad). 179 ANNALS OF THE CARNEGIE MUSEUM. This species is cited by Conrad from the ‘‘ Mahoning River, near Pittsburgh,’’ while Lea says: ‘* Mahoning River, Ohio.’’ The point of the Mahoning River nearest to Pittsburgh is where it joins the Shenango, thus forming the Beaver, near Mahoningtown, Lawrence County, about forty-five miles from Pittsburgh. The species is still present there, and I regard this part of the Mahoning River as the type-locanty. Unio viridis Conrad (Monogr., 4, 1836, and U. tappanianus Lea, Observ., II, 1838) = Symphynota viridis (Conrad). From the ‘‘ Juniata River,’’ according to Conrad, and from the ‘« Juniata at Hollidaysburg (Blair Co.)’’ according to Lea. Although Unionidee have recently been collected at Hollidaysburg, this species is not among them. Nevertheless it is present in the Raystown 3ranch of the Juniata River, and even farther west, in the head- waters of the West Branch of the Susquehanna in Indiana County. In 1891, E. H. Harn published a list of shells from western Penn- sylvania (Nautilus, 4, 1891, pp. 136-137). Harn was located at Blairsville, Indiana County, and apparently most of his Unionidze were from the Kiskiminetas. or the Conemaugh drainage. It is much to be regretted that this list does not give exact localities, since at present the fauna of the Kiskiminetas and its tributaries is almost com- pletely destroyed, only meager remains surviving in a few of the head- waters. Harn’s list is here submitted. All the species have recently been found in western Pennsylvania. The correct name according to the modern nomenclature is given. 1. Unio esopus Green = Pleurobema esopus (Green). 2. ‘¢ alatus Say = Proptera alata (Say). ‘© circulus Lea = Odovaria circulus (Lea). ‘© clavus Lamarck = Plewrobema clava (Lamarck). crassidens Lam. = Unio crassidens Lamarck. ‘© cylindricus Say = Quadrula cylindrica (Say). ‘¢ =gibbosus Barnes = Unio gtbbosus Barnes. 8. ‘ fabalis Lea= Micromya fabalis (Lea). O.2 “Onis der Lampriis.472s- lees to. ** kirtlandianus Lea = Quadrula subrotunda kirtlandiana (Lea). 11. ‘* Jligamentinus Lamarck = Lampsilis ligamentina (Lamarck). I Aum &: W| ORTMANN: UNIONTID2 OF WESTERN PENNSYLVANIA. 180 12. Unio multiradiatus Lea = Lampsilis multiradiata (Lea). ‘¢ mytiloides Rafinesque = Quadrula obliqua (Lamarck) Da. (form: pyramidata (Lea) ). ‘6 obliquus Lamarck = Quadriula obliqua (Lamarck). 14. 6 occidens Lea = Lampsilis ventricosa (Barnes). 15. ‘** parvus Barnes = Lampsilis parva (Barnes). 16. ‘*¢ phaseolus Hildreth = Ptychobranchus phaseolus (Hildreth). 17. ‘¢ pustulosus Lea = Quadrula pustulosa (Lea). 18. ‘* rectus Lamarck = Lampsilis recta (Lamarck). Ig. ‘** securis Lea = Plagiola securis (Lea). ‘6 ~subovatus Lea = Lampsilis ventricosa (Barnes), see no. 14. 20. §¢ subrotundus Lea = Quadrula subrotunda (Lea). 21. Anodonta undulata Say = Strophitus undulatus (Say) (incl. edentulus (Say) ). 22. Margaritana marginata Say = Alasmidonta marginata Say. 2)3) oe rugosa Barnes =.Symphynota costata ( Rafinesque ). 24. Ch undulata Say. = Alasmidonta undulata (Say). If the last species is correctly identified by Harn, it cannot be from the Kiskiminetas drainage, since it is positively absent from the Ohio drainage. Yet the species is found not very far from Blairsville, in the drainage of the West Branch of the Susquehanna (Clearfield Creek) in Cambria County. Of this list of twenty-three species (disregarding the eastern form just mentioned), nine have been found recently in the Kiskiminetas drainage, although of some of them only dead shells have been secured, or seen; viz.: Lampsilis ventricosa, L. multiradiata, L. lgamentina, L. recta, Ptychobranchus phaseolus, Strophitus undulatus, Symphynota costata, Alasmidonta marginata, Unio gibbosus, and Pleurobema clava. Two additional species not enumerated by Harn have been recently found in the Kiskiminetas drainage: Anodonta grandis Say and Quadrula coccinea (Conrad). ‘The rest of the fauna is irreparably lost in the Kiskiminetas drain- age, but most of these species still exist in the Allegheny River in Armstrong County. In order to now find Zampsilis parva and Quad- rula kirtlandiana, we are compelled to go up the Allegheny River as far as French Creek and its tributaries. The absence of Lampszlis luteola (Lamarck) in Harn’s list is re- markable. But this species, which is common in other parts, is also at present absent from the Kiskiminetas drainage, and this makes it 181 ANNALS OF THE CARNEGIE MUSEUM. the more probable that Harn’s shells were distinctly from this drain- age system. ; W. B. Marshall published (Bulletin New York Museum, 1, 1892, pp. 3 e¢ seg.) a list of Unionidee from New York. ‘The species were ‘ edvugtiments NOTES UPON THE FAMILIES AND GENERA OF THE NAJADES. By ARNOLD E. ORTMANN, PH.D. (Issued July 16, 1912.) d {Reprinted from ANNALS OF THE CARNEGIE MusEuM, VOL. VIII, No. 2, 1912] NOTES UPON THE FAMILIES AND GENERA OF THE NAJADES. By ARNOLD E. ORTMANN, Pu.D. Reprinted from ANNALS OF THE CARNEGIE MuseuM, Vol. VIII, No. 2, 1912.1 X. NOTES UPON THE FAMILIES AND GENERA OF THE NAJADES. By ARNOLD E. ORTMANN, PH.D. (Plates XVIII-XX.) In accordance with the observations recorded in a number of shorter notes on the Najades published recently by the writer (Ortmann, 1910a, I910b, 1910c, 1910d, 1911a) it is evident that the system of Simpson (19000) should be thoroughly revised, and that the soft parts of every species of mussels should be studied. In preparation for a monograph of the Najades of Pennsylvania this has been done by the writer, and the general results have been recently published in the first part of this work (Ortmann, 1911). But since it will take some time before the subsequent parts, dealing with the single species, will be ready for publication, and since the writer has examined, in addition, a great number of species not found in Pennsylvania, it seems well to publish these results as early as possible, combining the same with an attempt to rearrange the system to suit the new points of view. In the present paper, a general synopsis of the system will be given, in it assigning to each species, which has been examined, its proper place. Remarks as to the Figures.—For the majority of the genera, text- figures have been introduced to illustrate their principal characters. If possible, the type-species has been selected. These figures have been drawn from actual specimens, and are about natural size, but they have been generalized and are of a diagrammatic character, the chief features being emphasized. This refers chiefly to the gill-fila- ments (where they are given, as in the Margaritanide@) and the septa. The latter always are heavier than in nature, to bring out their charac- teristic features. In all figures the lettering is uniform, and the letters have the following meaning: an = anal opening; o = outer gill; br = branchial opening; pb = pes (foot); f =flapsof margin of mantle; pp = papille on margin of mantle; 1 = inner gill; sa@ = supra-anal opening. mp = marsupium; 222 ORTMANN: FAMILIES AND GENERA OF NAJADES. 223 The main figure always represents the soft parts seen from the left side, with the left half of the mantle removed. The Najades have been divided into three families: Margaritanide, Unionide, Mutelide. The first is holarctic; the second is known from Eurasia and North America, but probably exists also in Africa; the third is restricted to Africa and South America.! Family I. MARGARITANIDZ Ortmann. Diaphragm incomplete, formed only by the gills: outer lamina of outer gills only in part connected with the mantle, posteriorly free for a considerable distance. Anterior end of inner gills separated from the palpi by a wide gap. The margins of the mantle do not unite or approach each other anywhere, and there is no tendency to form branchial and anal siphons, and no supra-anal opening is present. Gills without water-tubes, interlamellar connections irregularly scat- tered, or forming irregular, oblique rows, or incomplete septa, which run obliquely to the direction of the gill-filaments. Marsupium formed by all four gills. Glochidia small, semicircular and globular, without hooks, but with irregular small teeth at the ventral margin. Family II. Unronrp# Swainson (restricted). Diaphragm complete, formed only by the gills: the outer lamina of the outer gills connected with the mantle at its posterior end. An- terior end of inner gills separated from the palpi by a more or less wide gap. Margins of the mantle drawn together by the gill-diaphragm, but not united, thus separating the anal from the branchial opening, and the anal is generally closed above by the union of the margins of the mantle (it rarely remains open), and, when closed, it always leaves a supra-anal opening (which is very rarely obliterated). Gills always with water-tubes, formed by interlamellar connections de- veloped as continuous septa, running parallel to the gill-filaments. Marsupium formed by all four gills, or by the outer gills alone, or by parts of the outer gills. Glochidia of various shapes, suboval, subtriangular, or celt-shaped, with or without hooks on the ventral. margin. 1The writer is convinced that the Najades will prove to be a most important group for the reconstruction of the ancient geographical features of the earth. As long as our knowledge of the systematic relations was obscure, or even directly wrong, any attempt in this direction must have been a failure. 224. ANNALS OF THE CARNEGIE MUSEUM. This family is divided into three subfamilies, as follows: 1. Subfamily UNIonIN2 Ortmann. Rarely no supra-anal opening formed, it is generally present, sep- arated from the anal opening by a shorter or moderately long mantle- connection. Marsupium formed by all four gills or by the two outer ones, when charged, only moderately swollen, and its edge not dis- tending. No secondary water-tubes developed within the marsupium. Glochidia rather small, or of medium size, subovate, without hooks; or subtriangular, with hooks. 2. Subfamily ANODONTIN2 Ortmann. Supra-anal opening always well separated from the anal opening, often by a very long mantle-connection. Marsupium formed only by the two outer gills, when charged, greatly swollen, and an extra thickness of tissue at the edge permitting the gills to distend. Within the marsupial gill, the water-tubes are divided during the breeding season into two lateral (secondary) water-tubes lying toward each face of the gill, and a central ovisac, which is closed at the base of the marsupium. Glochidia rather large, subtriangular, with hooks. 3. Subfamily LAMPSILIN® Ortmann. Supra-anal opening always separated from the anal opening by a mantle-connection of medium length, rarely entirely closed. Mar- supium formed by the two outer gills, or by parts of the latter, gener- ally situated in their posterior portion. When charged, the marsupium extends beyond the original edge of the gill, an extra thickness of tissue at the edge permitting a bulging out. Water-tubes of mar- supium not subdivided, when charged. Often special structures (papille or flaps) on the edge of the mantle, chiefly of the female, in front of the branchial opening. Glochidia of various sizes and shapes, small to rather large, either subovate, without hooks, or celt-shaped, with two spines on each valve. Family III]. MutTrE.ip# Gray (emended).? Diaphragm complete, formed anteriorly by the gills, posteriorly by a firm union of the margins of the mantle. Anterior end of inner gills * As has been stated in a previous publication (Ortmann, ro11a), the nomencla- ture of this family and its subfamilies is only provisional, until additional genera (chiefly Mutela itself) have been investigated. ORTMANN: FAMILIES AND GENERA OF NAJADES. 226 in contact with the palpi. Branchial and anal siphon sharply sepa- rated by the union of the margins of the mantle. Anal opening open, or closed above, in the latter case without forming a supra-anal open- ing. In some genera the margins of the mantle unite also in front of the branchial opening. Gills with very indistinct intercommunicating water-tubes, and interrupted interlamellar connections; or with well- developed water-tubes and septa, parallel to the filaments. Mar- supium formed only by the inner gills. The Jarve are glochidia or lasidia. This family is divided into two subfamilies. 1. Subfamily HyriiNn# Ortmann. Anal opening closed above. Marsupium with septa-like, inter- rupted, interlamellar connections, forming incomplete, communicating water-tubes. Non-marsupial gills with poorly developed interlamellar connections. Larva a glochidium. 2. Subfamily MUTELIN2 Ortmann. Anal opening open or closed. Marsupium with well-developed, continuous septa, forming well-defined water-tubes; also non-marsupial gills with septa and water-tubes. Larva a lasidium (?). There is no doubt, that of these three families that of the Margari- tanide is the most ancient; the lack of any tendency to form siphons, the incomplete diaphragm, the absence of real septa and water-tubes in the gills, the absence of a division of function in the gills (all four gills in the female are used both for breathing and for receiving the eggs), are peculiarities, which establish the primitive character of this family. The forward step in the development of the Unionide@ consists chiefly in the specialization of the marsupial structure. In the most primitive forms, all four gills are used as marsupia, but later on a division of labor is effected, so that in the female some gills serve only the purpose of respiration, while others, or parts of them, become organs used in propagation. But always, in this family, the inner structure of the gills is more complex than in the Margaritanide, which is expressed, by the development of septa and water canals;? 3In Margaritana monodonta a slight tendency is shown to develop septa, but here the septa are entirely different from those of the Unionidae, not running parallel to the gill-filaments, as in the latter, but diagonally to them. 226 ANNALS OF THE CARNEGIE MUSEUM. and secondly by the fact that the structure of the marsupium in the Unionide becomes highly specialized, or, to express it concisely, this family makes a special effort to bring the marsupial apparatus to the highest degree of efficiency. All differentiation is connected with two purposes: the lengthening of the breeding season, and the change of the discharge of the glochidia from a ‘“‘natural”’ to an “un- natural’’ manner, if such an expression may be allowed. As regards the first, the subfamily of the Unionine is as yet in an undifferenti- ated condition, possessing a short breeding season (being tachytictic), without specialization.4 But in the Anodonting and Lampsiline the breeding season is extended over the winter and the glochidia, after they are fully developed, are not discharged immediately, but retained for a long period in the marsupium (bradytictic). This renders it necessary to develop special devices in the marsupium, and the most urgent need apparently is to provide the necessary oxygen for the glochidia enclosed in the marsupia. It is now interesting to observe how this purpose is accomplished in two different ways by the two subfamilies. In the Anodontine, the lateral, secondary water-tubes cut off from the central ovisac, undoubtedly have the purpose of keeping up a lively current of water around the swollen marsupial mass. Nothing similar to this is known in the Lampsiline, but in the case of these the whole marsupium bulges out beyond the original edge of the gill, and this bulging mass is enclosed in a rather thin membrane, favoring osmotic processes. Further, there is a tendency to locate the marsupium in the posterior part of the gill, and to push it toward the lower posterior end of the shell, so that it is close to the branchial opening, where fresh and pure water enters the animal. In addition, a number of the Lampsiline develop special papille and flaps on the edge of the mantle, just at the place toward which the marsupium is pushed, and these structures surely have the purpose of producing a lively current of water over the marsupium. Further- more, the thin membrane enveloping the protruding part of the marsupium, and its position near the branchial opening, are apparently connected with the peculiar discharge oi the glochidia in the Lamp- siline, which is through the edge of the marsupium, through holes 4 Haas (1910e, p. 19) comparing the marsupium of Anodonta and Unio expresses the opinion that that of Anodonta is more primitive than that of Unio, which is entirely erroneous: just the opposite is the case, that.of Anodonta being much more complex and specialized. As long as views like this prevail, we cannot expect to arrive at a proper understanding of the system of the Najades. ORTMANN: FAMILIES AND GENERA OF NAJADES. 227 which form there for this purpose. This ‘“‘unnatural’’ discharge is known only in the Lampsiline, and is unknown in the Anodontine and Unionine. Finally the family of the Unionide differs from the Margaritanide and is more highly advanced in the formation of rudi- mentary siphons. But in this respect this family is not very progres- sive. It has the anal and branchial openings separated only by the (complete) gill-diaphragm, and in addition, it has the anal closed above, thus giving it an incomplete tubular shape. Beyond this, there is no progress in this family. The presence of a supra-anal opening is, in my opinion, only incidental to the closing of the anal. The members of the third family, the Mutelid@, have gone in another direction in their development. If the expression may be permitted, they lay chief stress upon the better development of the siphons, while in the differentiation of the gill-functions they have started out from the beginning with another idea, which, however, has not attained a very high degree of perfection. With regard to this it may be said that they have restricted the marsupial function to the inner gills, and very likely the anterior connection of these gills with the palpi is incidental to this function. Not much advance is to be observed in the gill structure, and only two types are met: incomplete septa and intercommunicating water-tubes (a rather primitive condition) in one group (Hyriine); and complete septa and water-tubes in the other group (Muteline). Very likely the latter structures are not homolo- gous to the septa and water-tubes of the Unionide, but have been ac- quired independently, since their finer structure is different. With regard to the siphons, which attain within this family their highest perfection among the Najades, we have first of all a complete separa- tion of anal and branchial openings by a firm mantle-connection, which forms the posterior continuation of the gill-diaphragm, and in addition we have a tendency to close both the anal above, and the branchial below, by mantle connections. It is true that this tendency is not yet perfect in many Mutelide, but it is developed within this family, so that in the most highly specialized genera we have two real: tubular siphons, formed by complete coalescence of the edges of the mantle. I think the above account of the phylogenetic tendencies within the various divisions of the Najades will make it clear that the mor- phological characters upon which our new system is founded are 228 ANNALS OF THE CARNEGIE MUSEUM. characters which are essential, since they indicate the various ‘‘ideas”’ in the specialization within each group, and advance our understanding of the phylogenetic progress and the systematic affinities of the Na- jades. It may not be amiss to point out that it is absolutely impossible to recognize this system in the characters of the hard parts, the shells. It is true that certain types of shell are characteristic within smaller groups, and that there are cases, where we are able to recognize a genus, for instance, by the shape of the shell. But if we come to compare the subfamilies and families, we find that various types of shell turn up in them again and again. This goes so far that certain species resemble each other so much externally that they have been confused or placed together even by our greatest authorities, while they actually may belong to entirely different groups according to the soft parts. For this reason I have deliberately omitted to give shell characters for the families and subfamilies, for this is simply impossible. One character of the shells, however, may be of greater value, and this is the beak-sculpture. As will be seen below, I shall use it re- peatedly for the definition of genera. But it has been largely misunder- stood, and is even now not very clear. Simpson, in distinguishing a concentric and a radial beak-sculpture, made a great mistake in uniting under the latter two types of sculpture, the radial and the zig-zag, while he united the double-looped with the concentric sculpture. According to my studies, which, however, are not yet fully satis- factory, the following seem to be the real conditions: The original and simplest beak-sculpture consists of concentric bars. A few (one to two) of them are, when the beaks are well preserved, always present, even in zig-zag or radially sculptured beaks. In many forms other bars of the same character are added, and no complications are ob- served. In other forms the later bars become double-looped. This character is generally inaugurated by the fact that the posterior part of the simple bar, which lies upon the posterior ridge of the shell, is emphasized. It becomes more pronounced, often tuberculiform, and is drawn out in the direction of the posterior ridge, toward the lower posterior angle of the shell. This produces an angular projection in the posterior part of the original bar, which by contrast with the anterior part, which does not project, gives the appearance of the bar consisting of two parts, or two loops, till we finally come to a beak- sculpture which distinctly consists of a double loop, the two parts ORTMANN: FAMILIES AND GENERA OF NAJADES. 229 separated by a distinct reéntering angle. In some forms these two parts become tuberculiform, and the connecting bars disappear, so that the beak sculpture appears composed of isolated tubercles (Unio pictorum). A further step in advance is that the anterior part of the double-looped bar breaks up into tubercles, and finally into zig-zag bars. The manner in which this is accomplished remains yet to be studied, but always, in the zig-zag sculpture, the posterior loop, which lies upon the posterior ridge, is the most conspicuous part of the whole beak sculpture. Thus the zig-zag sculpture is the most extreme con- dition of a line of development, which goes from the simple concentric bar, through the double-looped, to the zig-zag condition. In certain forms with zig-zag sculpture, all three stages are clearly present on the same shell (Nodularia douglasie), and possibly this may be observed always in such cases.° Entirely different from this is the radial sculpture, but I have reason to believe that it also goes back to the concentric type. At any rate, I have seen in specimens of the genus Lamellidens that there are also originally one to two simple concentric bars. But after these only the lateral parts of the bars, which anteriorly and posteriorly curve up toward the beak, are developed, while the middle part upon the disk becomes obliterated. Then these lateral parts, which have a direction from the beak toward the basal margin, are emphasized, developing more strongly, and their direction remaining a radial one. Since there are two groups of radiating ridges (an anterior and a posterior), the median ones naturally must interfere with each other upon the middle of the disk, and must come in contact there at a more or less sharp angle, when fully developed. This is in fact the case, wherever we see radial sculpture well developed. There are always two sets of radiating folds or ridges, one originating in front, the other behind the beak, which cover the disk interfering with each other in the middle of the shell. In some cases, however, this is not very clear, and such cases possibly present the highest type, with the original features obliterated. The radial sculpture is another extreme standing at the end of a line of development starting from simple concentric loops. These conditions are worthy of being studied more closely. In regard 5 In certain cases it seems that double-looped sculpture may again be simplified by the re-entering angles becoming less sharp and only sinuate in the latest bars. But this is surely a sign of the incipient obliteration of sculpture. 230 ANNALS OF THE CARNEGIE MUSEUM. to its systematic value, the beak-sculpture, when properly understood, indicates certainly systematic affinity, but is not fit to be used for the distinction of larger groups, since it is very likely that the different types were developed rather early, and are found side by side among the more primitive groups of Najades, the Unionine for instance. Yet in the more advanced groups often only one type isfound. Thus, for instance, among the Anodontine and Lampsiline, we possess only the first type up to the double-looped structure, while the zig-zag structure is practically absent, and no trace of the other type (the radial) is ever found. On the other hand, it seems that in the Mu- telide only the radial type is present, provided there is any sculpture at all. In addition, conditions become yet more complex by the fact that the beak-sculpture in general seems to be a character which is subject to obliteration, and anywhere within the system we may expect to meet forms which have reduced their beak-sculpture to a lesser - or greater degree, often to complete disappearance. ‘Thus we may say, in a general way, that beak-sculpture, although important and in- dicating the minor affinities, is unfit to be used for the distinction of the larger groups. Family MARGARITANID~. I recognize only one genus in this family, to which a number of species have been assigned by Simpson, of which, however, the structure of four only is known. Genus MARGARITANA Schumacher, 1817. Simpson, 1900), p. 674. Margaritana margaritifera (Linnzus). Some twenty specimens are at hand, from the drainage of the upper Little Schuylkill River in Schuylkill County, Pennsylvania, collected by myself; soft parts of another specimen from the Auma creek, near Weida, Saxe-Weimar, Germany (drainage of Elster river); and three complete specimens from the Perl-Bach at Postfelden, near Falken- stein, Bavarian Forest, Germany. For these German specimens I am indebted to Mr. W. Israél. Published figures: Photograph of soft parts in shell, by Carl (1910, pl. 4, figs. A and B); of gills, by Ortmann (19110, p. 285, fig. 1, and pl. 87; fig: 11). ORTMANN: FAMILIES AND GENERA OF NAJADES. 231 I have found that the German specimens agree in every particular with the American form. Certain characters not observed in my specimens (marsupium and glochidia) I have gathered from the literature (chiefly Harms, 1907 and 1909). Margins of mantle free all around from the anterior to the posterior end and with no tendency to unite anywhere. Branchial opening indistinctly separated from the anal, a horizontal ridge running from the posterior insertion of the outer lamina of the outer gill to the margin of the mantle, but the margins of the mantle are not held together by the diaphragm. Anal opening not closed above, and no supra-anal Fic. 1. Margaritana margariltifera (Linneus). Specimen from the Perl-Bach, Postfelden, Bavarian Forest, Germany. (Carn. Mus., No. 61, 4,987.) formed. Branchial opening ill-defined anteriorly, on the inner edge with strong papillae, which disappear anteriorly, and then the inner edge of the mantle is smooth. Inner edge of anal opening almost smooth. Palpi large, subfalciform, drawn out and pointed behind, their posterior margins united for about one-half to three-fourths of their length. Gills long and broad, the inner the wider, chiefly so anteriorly. Outer gill becoming gradually narrower in front, its anterior end at the highest point of the line of attachment of the mantle, high above the palpi. Inner gill narrowing more suddenly anteriorly. Its anterior end is found in front and below that of the outer gill, but separated from the palpi by a wide gap.® Edge of inner gills with a longitudinal furrow, which is absent in the outer gills (this character is present in all Najades examined, and will not be mentioned again). 6 This character is variable, and sometimes (as in the specimen which served as: original for our figure) the gap is rather short. 2ae ANNALS OF THE CARNEGIE MUSEUM. Outer lamina of outer gills not entirely connected with the mantle, but its posterior part is free for about one-fourth, or slightly less, of its length. Inner lamina of inner gill free from the abdominal sac with exception of its anterior end. Behind the foot, the two inner lamine of the inner gills are connected. Thus the diaphragm (separation of branchial and suprabranchial-cloacal cavities) is formed only by the gills, and it is incomplete posteriorly, and does not reach the margin of the mantle, although the medially united free ends of the gills project to near the margin of the mantle. Gills without water-tubes and without septa. The interlaminar connections are patch-like, irregular in shape and position, and only here and there a diagonal arrangement, from the base toward the edge and forward, is indicated, which, however, does not follow the direction of the gill-filaments, and does not form continuous septa. Color of soft parts grayish, inclining to blackish. Foot brown, paler on edge; through the middle runs a black band, sharply marked off from the whitish abdomen, but gradually shading into the brown of the foot. Gills brownish-gray anteriorly, shading to black pos- teriorly. Mantle brownish-white, edge black, broadly so behind. From the posterior end of the attached part of the outer lamina of the outer gill there runs to the posterior margin of the mantle a white line, bordered below with black. No gravid females are at hand, and thus the writer cannot say any- thing about the marsupium and the glochidia. However, these have been described by others (see Harms, 1907 and 1909). The marsupium is formed by all four gills,?7 and the glochidia are very small (0.0475 mm.), semicircular, globular, without true hooks, but with a number of small teeth on the ventral margin. The breeding season in Pennsylvania is in June and August (Conner, 1909, p. 112), in Germany in July and August (Harms, 1907, p. 814), and probably twice in succession during this time (Harms, 1909, p. 332). Margaritana sinuata (Lamarck). The anatomical structure of this species has been described by Haas (1910), p. 181), who created for it the new genus Pseudunio. Although * Simpson (1900), p. 674) says that the marsupium is formed only by the outer gills, and refers (footnote 2) to von Wahl. I have consulted this paper (Wahl, 1855), but cannot find in it anywhere a description or mention of the marsupium of Margaritana. ORTMANN: FAMILIES AND GENERA OF NAJADES. 233 Haas omitted to say anything about the gill-structure, his description of the margins of the mantle, of the diaphragm, and other parts renders it absolutely certain that this species must be placed in the family Margaritanide. Haas points out certain differences from Mar- garitana margaritifera, of which the most important is the fact that the shell has lateral hinge-teeth. Since we have other genera among the Najades in which the hinge-teeth are variously developed, and since it is absolutely clear, that Margaritana margaritifera, without lateral teeth, must have descended from forms with such teeth, I think the differences in Unio sinuatus should be regarded as only of specific value, and I see no reason why we should not place it with Margaritana, with which some of its most essential and important characters are known to agree, while all the known differences are such as in other groups are known to be of minor value. Margaritana sinuata thus would represent a somewhat more ancient type than M. margaritifera (see Ortmann, I9II¢, p. 6). Margaritana monodonta (Say). I have received, from B. Walker, one complete specimen, and the soft parts of three others, all from the Cumberland River in Pulaski, Russell, and Cumberland Counties, Kentucky. Fic. 2. Margaritana monodonta (Say). Specimen from Cumberland River, Rowena, Russell Co., Ky. (Carn. Mus., No. 61, 4,960.) We may compare the description of the soft parts by Lea (Obs., X, 1863, p. 422), which, however, mentions among the important features only the posterior end of the gills, the branchial and anal openings. Margins of the mantle, branchial and anal openings as in M. mar- garitifera. No supra-anal present. Posterior margins of palpi connected for about one-third of their length. Gills rather long (cor- responding to shape of shell), the inner the wider, chiefly so ante- 234 ANNALS OF THE CARNEGIE MUSEUM. riorly. Anterior ends of the two gills as in M. margaritifera. Outer lamina of outer gill not entirely connected with the mantle: posteriorly a part of it is free (about one-seventh of length of gill, but probably more, since this part in all my specimens is considerably contracted). Inner lamina of inner gill asin M. margaritifera, and thus the structure of the diaphragm is essentially the same. Both gills are quite delicate. The two lamine are not connected by septa running parallel to the gill-filaments, but the inter- laminar tissue forms septa of another type: they run obliquely, diagonally, from the base of the gill downward and forward. Of these septa some are longer, others shorter, and toward the edge of the gill they sometimes curve a little in the direction of the filaments. The septa thus are rather irregular, and being quite distant from each other, no regular water-tubes are formed. A difference in the arrangement of the septa, which might be due to sex, could not be observed in the specimens at hand. None of them was gravid, so that nothing can be said wither anacimen’ from about the arrangement of the ova in the gills, same locality. and about the glochidia. The fact, that in all specimens the structure of the two gills is practically identical, suggests, however, that all four gills are used as Fic. 2a. Left gills of marsupia. Color of soft parts whitish, edge of mantle blackish all around, but chiefly at the anal and branchial openings. Gills transparent, but not blackish. Foot grayish-white in its distal part, this gray color marked off in a sharp line from the basal white part. M. monodonta agrees in most characters with M. margaritifera, and chiefly in the general form of the margin of the mantle, the branchial and anal openings, the diaphragm, and the structure of the gills. The chief difference is found in the diagonal, incomplete septa of the gills, which, however, unmistakably correspond to the irregular diagonal rows of interlaminar connections in M. margaritifera. In this respect, M. monodonta represents a stage of development slightly more in advance of that of M. margaritifera, and this would support the view ORTMANN: FAMILIES AND GENERA OF NAJADES. 235 expressed by Walker (19102, p. 137) that it is an “offshoot of the more ancient margaritifera-stock.’’ In its general appearance it looks rather like a depauperated form, while the gill-structure has attained a slightly higher stage of differentiation. It also differs slightly in the lesser development of the papilla of the branchial opening, and the great reduction of the black color suffusing most of the soft parts of M. margaritifera, and, of course, also in shell characters. If we should accept the genus Pseudunio proposed by Haas for U. sinuatus, we would have, as a simple logical consequence, to create a new genus for M. monodonta, for the difference of the gill-structure of the latter is much more important than any of the differences knownin M. sinuata. It represents a phylogenetic step in advance. But having to deal only with four species, I see no reason why we should not leave them together in the genus Margaritana. Margaritana hembeli (Conrad). Eighteen specimens from Hunters Creek, Evergreen, Conecuh Co., Alabama, have been investigated. They were collected by H. H. Smith on February 10, 1911. Fic. 3. Margaritana hembeli (Conrad). Specimen from Hunters Creek, Ever- green, Conecuh Co., Ala. (Carn. Mus. No. 61, 5,022.) Margins of mantle, branchial and anal openings, and gills much as in M. margaritifera. Papille of branchial rather small. Posterior mar- gins of palpi connected for about three-fourths of their length. Interlaminar connections of gills irregularly scattered, here and there with a tendency to fall into oblique, irregular rows. On the whole this tendency is less developed than in M. margaritifera, but there is some variation in this respect in different individuals. I was unable to dis- 236. ANNALS OF THE CARNEGIE MUSEUM. cover any marked differences in the structure of the gills which might be due to sex. No gravid females were found. Color of soft parts brownish-white; foot grayish-brown, the darker part sud- denly marked off in a sharp line from the white abdominal sac; palpi and gills brownish, the latter more grayish pos- teriorly; mantle pale brown, its margin whitish with brown-black edge, most intense posteriorly; a black line on mantle separating anal and branchial cavities. This is a true Margaritana, much resembling in structure WM. margaritifera. But it has well-developed lateral hinge- teeth, and thus must be considered as a more primitive type. It has no closer relationship with M. monodonta, and iG) Ga Glefteills tof another) SCemMOn be connected with it. In shell- specimen from same locality. sculpture, M. hembeli is quite unique. Its distribution (in southern Alabama and Louisiana) offers a very interesting problem. Family UNIONID-. Subfamily UNIONINZ. Simpson's (1900b) North America genera: Quadrula, Tritogonia, Pleurobema, and Unio belong to this subfamily. Further, I have shown (Ortmann, r9IIc) that the European Unio also belongs here, as well as the Asiatic genera Parreysia and Lamellidens. I have further demonstrated, that the European Unio is not identical with the North American Unio, and that for the latter the generic name of Elliptio should be used. The genus Tyitogonia is simply a synonym of Quadrula in Simpson’s sense (see Sterki, 1907, p. 48, and Ortmann, I9IID, p. 329). In the structure of the soft parts there is not much differentiation in all these forms. The most important is that in some all four gills are used as marsupia (see Plate XVIII, fig. 1), in others only the two outer ones (see Plate XVIII, figs. 2, 4,5). The systematic value of this character has been doubted (see Frierson, 1909, p. 107). Yet I believe ORTMANN: FAMILIES AND GENERA OF NAJADES. 237 that it is of prime significance. In all my investigations I have never come across an exception or variation inthe marsupium. It is true that in species which have normally all four gills marsupial, sometimes only two gills are found charged. But all authors, who record such cases, only mention the fact that the gills were charged, without saying any- thing about the structure of the gills. I have also met with such cases: but invariably a closer investigation revealed the fact that the other gills which were not charged also possessed marsupial structure, and consequently were capable of being charged with eggs. On the other hand, in those cases, where the outer gills alone serve as marsupium, it was not the simple fact that they alone were filled with eggs in the breeding season, which was ascertained. It was the in- vestigation of the structure of the gills, which induced me to judge the character of the marsupium. Lefevre and Curtis (1910, p. 83) are inclined to regard my observations in Pleurobema coccineum as due to accidental conditions. But this is surely not so. I have seen now a great number of individuals of this species with the outer gills alone charged, and I have seen and examined many more, females in the sterile condition, which invariably had marsupial structure only in the outer gills, while the inner gills were different, and not built to receive eggs. Not a single exception was observed. Further it is quite evident that the arrangement of four gills serving as marsupia is found in a number of groups, the species of which are undoubtedly closely allied. This is clear in the peculiar Quadrula piicata-group, in the metanevra-group and others. Then again, a marsupium formed by the outer gills alone is characteristic of other natural groups. To me the most interesting case was that of Pleuro- bema coccineum. Here I discovered first that this supposed Quadrula differs from the Quadrula-type; I also discovered that this species intergrades with Q. obliqua and Q. pyramidata. This being the case, I concluded that the latter also should have a marsupium like coc- cinea. And this proved to be true! Nevertheless the character of the marsupium should not be too implicitly relied upon. There is no question that the condition in which the four gills serve as marsupia is more primitive than the stage where only the outer gills are marsupial.8 But it seems to me that 8 The functional and morphological progress from the four-gill-marsupium to the two-gill-marsupium has been correctly understood and expressed by Haas (1910e, p- 19). 238 ANNALS OF THE CARNEGIE MUSEUM. there is or has been a general tendency to restrict the marsupium to the outer gills, and that this forward step in a quite natural direction has been made independently in various groups. That is to say, the Unio-type of marsupium has repeatedly developed from the Quadrula- type by parallel evolution. Of the other features of the soft parts only three furnish some help for the distinction of genera. The first and most important is the character of the placenta, revealing differences which are of prime value, but affect only a few forms, as will be seen below. The second is the separation of the anal and supra-anal openings. Although characteristic of certain forms (in one case these openings are not at all separated), it is somewhat variable in others, even individually. Thus we can use this character only to a limited degree. The third is the connection of the inner lamina of the inner gills with the ab- dominal sac. Here there seems to be a difference between certain forms of the Old and the New World. But, unfortunately, too few of the former are known for me to express a final judgment. Thus the soft parts alone would furnish only few criteria for the distinction of genera, and we should direct our attention to the shell. Here we have indeed great variety, and the shapes of the shell have been largely used heretofore for the definition of genera. The most important feature, in my opinion, is the beak-sculpture, which, how- ever, has been largely misunderstood by Simpson. In fact in this primitive subfamily we have, side by side, all the different types of beak-sculpture, and, as we shall see, they may be used to great advantage. Since various types of shell-structure are frequently combined with various types of soft parts, it would not do to make only a few large generic divisions. For if we recognize, for instance, only two main genera according to the character of the marsupium, the same types of shell would turn up in either of them, which surely would give an incomplete or wrong impression of affinities. Thus, in my opinion, it is advisable to admit a larger number of genera founded upon both the structure of the soft parts as well as of the shells. Such a scheme is introduced here, at first, tentatively, but I hope it finally will prove to be the most convenient. Finally I should mention the glochidia of these forms. The latter are known in a number of North American.species, where they always are of a primitive shape (see Plate XIX, fig. 1). They are also ORTMANN: FAMILIES AND GENERA OF NAJADES. 239 known in European forms, where they incline toward the type of the subfamily Anodontine. I have no doubt that this finally will be a very important systematic criterion, but unfortunately we do not know the glochidia of a single Asiatic species. The following provisional division into genera in accordance with what has been hereinbefore said is here submitted: a. Beak-sculpture ranging from the concentric to the zig-zag type. Mantle con- nection between anal and supra-anal absent, deciduous, short, or of medium length. Inner lamina of inner gills free from abdominal sac. bi. Mantle connection absent or short. Beak-sculpture concentric to zig-zag. Glochidia subovate, without hooks. ca. All four gills serving as marsupia. Mantle connection between anal and supra-anal present, short and deciduous.’ d,. Ovisacs and placente subcylindrical, the latter rather persistent, generally red. Shell simple, without sculpture upon disk. Beak- Sculpture: sitnple, cOncemtricia. cs. ers. ei eieleta ee ns Fusconaja. d». Ovisacs and placentz leaf-shaped (compressed and lanceolate), the latter rather poorly developed, generally white. Shell with sculpture of various patterns. Beak-sculpture concentric, double-looped, or zig-zag. e1. Shell with oblique undulations upon the disk. Beak-sculpture concentric, nearly obliterated, or of zig-zag pattern and extending more or less upon the disk............ Crenodonta. eo. Shell-sculpture tuberculous or nodulose. Beak-sculpture con- centric and disappearing upon the disk, or of the double- looped or zig-zag pattern, more or less extending upon the disk. Quadrula. c.. Marsupium formed by the outer gills only. d;. Mantle-connection above anal opening absent, no supra-anal formed. Shell tuberculous, beak-sculpture of zig-zag pattern, much broken up, extending somewhat upon the disk. Nacre deep PUBP LS rey Morse cutee lest eheceeere seme CUS Inrtahe esa ke Rotundaria. dy. Mantle connection between anal and supra-anal present, short, or deciduous. Beak-sculpture concentric, obliterated toward the disc. é1. Shell tuberculous. Soft parts of a peculiar orange color. Pla- cente pink (at least in one species)........... Plethobasus. § Under this division apparently belong two species occurring in Georgia and Florida, infucata Conrad and kleiniana Lea, the soft parts of which have been partly described by Lea (Obs., X, 1863, pp. 404 and 407). Inthese species we observe the most beautifully developed zig-zag sculpture among North American forms. The soft parts are imperfectly known, but the marsupium is formed by all four gills. Probably they should form a genus by themselves. 10 Called ‘‘egg plates”’ by Lillie (1895), and ‘‘conglutinates’’ by Lefevre and Curtis {1Q10). 240 ANNALS OF THE CARNEGIE MUSEUM. ex. Shell without sculpture. Soft parts more or less whitish, rarely slightly colored. Placentz whitish, rarely slightly colored. fi. Shell elongate and oblique, with the beaks placed rather an- teriorly, or rounded, squarish, or even elevated. Epi- dermis light, brownish, rarely dark brown, with or without rays. Nacre light colored. Beak-sculpture concentric, poorly developed............. Pleurobema. fs. Shell more or less elongate, but not oblique, beaks not much anterior. Epidermis dark or light, generally without rays, or rays indistinct. gi. Shell with rather dark epidermis, sometimes faintly rayed. Nacre often dark (pink to purple). Beak- sculpture concentric, with an angle upon the posterior ridge, but not double-looped, often faint and rudi- MENtAry ce cytiad GRC EEA OE se Elliptio. g. Shell with lighter epidermis (often with dark bands), rays practically absent. Nacre whitish. Beak-sculpture rather distinct, concentric, bars not angled behind, but regularly Cunvedsiip.e sacl aemietaoe ence css: Uniomerus. bs. Mantle-connection between anal and supra-anal openings well developed, but generally shorter than the anal. Shell not sculptured upon the disk, elongated, but not oblique. Beak-sculpture sharply double-looped or of the zig-zag type. Glochidia subtriangular, with hooks.. .Unio. a2. Beak-sculpture of the radial pattern. Mantle connection between anal and supra-anal present, rather long. Inner lamina of inner gills connected with abdomincal sac. b;. All four gills marsupial. Beak sculpture radial, well developed, more or less ‘extendins-upon the jdisky.c...c.c thie a ons hs cree tie tee eae Parreysia. bs. Two outer gills only marsupial. Beak-sculpture concentric-radial, rudi- TIVELUUATNG, foes eetalae. set Sayers: ste eget nan Ae lae RA eae eat ae ee eee ate ee Lamellidens. Genus FUSCONAJA Simpson. 1900. Simpson, 1900), p. 784 (as section). I consider this the most primitive type of the Unionid@ known to me. Shell simple, rounded, ovate, quadrate, or triangular, with more or less elevated beaks, well developed hinge-teeth, and rather deep beak- cavities. Outer surface without sculpture. Epidermis lighter or darker brown, with hair-like, dark rays, sometimes fused into spots when young. Beak-sculpture simple, concentric, slightly angled upon the posterior ridge, but not double-looped, not extending upon the disk, and often obliterated. Soft parts of primitive structure. Supra-anal separated from the ORTMANN: FAMILIES AND GENERA OF NAJADES. 241 anal, but mantle-connection between them very short, and deciduous, often absent. Inner lamina of inner gills free from abdominal sac. All four gills marsupial. There is hardly any difference in structure between the inner and outer gill, When gravid, the water-tubes (ovisacs) do not expand much, and their lumen remains nearly cylin- drical. Placentze also subcylindrical, generally red in color, rather persistent, and discharged whole. Glochidia rather small, subovate, without hooks. Type: F. trigona (Lea), which (cf. Walker, 1910b, p. 24) should bear the name wndata (Barnes). Fusconaja undata (Barnes). About a half dozen specimens of the form from Lake Erie have been examined, and in July, 1910, I found a few gravid specimens. Mr. H. E. Wheeler sent two males, and six females (one of the latter gravid) from the Ouachita River, Arkadelphia, Arkansas, collected March 25) Lom, This form agrees in allessential points with F. rubiginosa. The ova, placente, and sexual glands have the same red color. The soft parts are less inclined to orange, are paler, and often whitish and cream- colored. Simpson (in Baker, 1898, p. 76) gives a rather meager de- scription. The glochidia are unknown, all specimens found by myself had only eggs. The gravid female from Arkadelphia was just beginning to charge the gills. This early date (March 21) should be noted. Fusconaja rubiginosa (Lea). Numerous specimens, in all conditions, have been examined, all collected in the smaller creeks of the Ohio drainage in western Penn- sylvania. This species is typically tachytictic, but the breeding season is rather long, from the middle of May to the beginning of August. In the case of single individuals it is probably much shorter. Descriptions of the soft parts have been given by Lea (Obs., X, 1863, p. 416) and Simpson (in Baker, 1898, p. 78). Edges of the mantle drawn together by the gill-diaphragm, thus separating the anal and branchial openings. Anal opening closed above by a very short mantle-connection, thus forming a very large supra-anal; but this mantle-connection is very inconstant and de- 242 ANNALS OF THE CARNEGIE MUSEUM. ciduous, often absent, sometimes torn. Branchial opening with papilla on inner edge, anal with distinct, but small papillae. Palpi subfalciform, pointed behind, their posterior margins connected for about one-third to one-half of their length. Fic. 4. Fusconaja rubiginosa (Lea). Male, from South Fork of Tenmile Creek, Waynesburg, Greene Co., Pa. (Carn. Mus., No. 61, 4,509.) _ Coll. May 6, 1910. Gills short and rather wide, with curved lower margins (correspond- ing to the shape of the shell), the inner gill wider. Outer gill attached at its anterior end at the highest point of the attachment-line of the mantle, far above the palpi; in- ner gill with its anterior end slightly in front and below that of the outer gill, widely separated from the palpi. Outer lamina of outer gills entirely connected with the mantle. Inner lamina of the inner gill free from the abdominal sac, except at its an- terior end. Behind the foot, the Fic. 4a. Left gills of a sterile female from : ; : two inner lamine of the inner same locality. gills are connected up to their posterior end. Thus a complete gill-diaphragm is formed, which reaches backward close to the posterior margin of the mantle. Both gills possess well developed septa and water-tubes, running parallel to the gill-filaments. Inthe male, the septa are rather distant and the water-tubes are wide. The septa are merely lines of con- nection of the interlaminar tissue. In the female, all four gills are marsupial, the septa are better developed, thicker and longer (in the ORTMANN: FAMILIES AND GENERA OF NAJADES. 243 transverse direction), more independent structures, with an epithelium thrown up into folds. They are much closer together, and form much narrower water-tubes, which, when gravid, become ovisacs. There is no noticeable difference in the width of these water-tubes in the inner and outer gill. When charged, the ovisacs do not expand much, and their lumen remains subcylindrical, so that the whole marsupium does not swell to any considerable degree, and its edge does not dis- tend and remains sharp. The ova are red in color, and are lodged in the ovisacs in the shape of well developed placente (sticking together by their membranes). The placenta, conforming to the shape of the ovisacs, are subcylin- drical, and are discharged whole through the anal opening. The glochidia (see Ortmann, 1911b, pl. 89, fig. 2) are rather small, of suboval shape, without hooks. Length and height about equal, 0.15 mm. The color of the soft parts is somewhat variable, but generally a yellowish-orange. The margin of the mantle, the distal part of the foot, and the adductor muscles, are deeper in color (intense orange- brown), while the gills are pale yellowish or brownish. The gills of the gravid female appear red when charged with the ova, and inside of the whitish abdominal sac the gonads are very oftenred. Incertain specimens the color of the soft parts is altogether paler, the bright orange tints being missing, but this difference in color does not depend on sex. Fusconaja cerina (Conrad). One male specimen at hand, received from L. S. Frierson. It is from Bayou Pierre, De Soto Parish, Louisiana. Structure in all points like that of F. rubiginosa, and agreeing also in minor details, such as the papille of the anal and branchial openings, separation of anal and supra-anal, inner lamine of the inner gills, and palpi. A female was not at hand. But Mr. Frierson writes to me con- ’ cerning this species: ‘‘eggs in four gills,’ and ‘‘cerina has the body white in about half, but red in the other half of the specimens. Some have red eggs. But red eggs and red body are not correlated.” Thus it seems that this species stands very close to F. rubiginosa, a relationship, which has been assumed by others on the ground of the characters of the shell. 244 ANNALS OF THE CARNEGIE MUSEUM. Fusconaja lananensis (Frierson). Frierson (1901, p. 76) describes the soft parts as: ‘‘salmon-colored, scarlet when cut’’ (probably sexual glands). ‘‘Eggs carried in all four gills, very red.’”’ This, together with the general shape of the shell, renders it almost certain that this species should be placed here. Fusconaja subrotunda (Lea). I have investigated numerous individuals in all conditions collected by myself in the Ohio and Allegheny Rivers in western Pennsylvania, and some additional ones from the Ohio between Pittsburgh and Cincinnati. The breeding season falls in June and July. The soft parts have been described by Lea (Obs., X, 1863, p. 427). This species agrees well with F. rubiginosa, but special mention should be made of the anal opening, which has fine crenulations, and shows the same variability as regards the short mantle-connection separating it from the supra-anal. The structure of the gills (see Ortmann, 1911}, pl. 86, figs. 1-3) is essentially the same as in F. rubiginosa. The ova are generally red, but in rare instances they are pale pink or white. The placente are also subcylindrical, and are discharged whole. The glochidia are similar to those of F. rubiginosa, but slightly higher than long.. Length 0.13 mm.; height 0.15 mm. In the color of the soft parts, two types may be distinguished. Normally there is much orange color present, which is most intense (deep orange-red) on the foot, the mantle margins, and the adductors, while the rest, chiefly the gills, are more brownish. When charged the gills are red. In the other type of color all parts are whitish or *yellowish, or brownish-white, the latter color chiefly on those parts, which are orange in the other type. Yet there are intergrades between these two types, the orange color gradually passing into the brown. In western Pennsylvania, the orange type prevails. There is no re- lation of these colors to sex. The color of the gonads deserves special mention in this species for comparison with the next. Inallspecimens examined it is either whitish or paler or deeper red, the latter of a distinct crimson hue, identical with the color of the eggs, but generally more intense. This crimson is found both in the male and in the female, so that it seems that also the sperm is thus colored. Sr ORTMANN: FAMILIES AND GENERA OF NAJADES. 24 Fusconaja ebena (Lea). Of this species I have examined only very few typical representa- tives, found by myself in the Ohio River at Portsmouth, Scioto County, Ohio (Sept., 1910). The soft parts of a gravid female have been figured by Lefevre and Curtis (1910, pl. 1, fig. 4). Although this figure is correct, the struc- ture of the gills is not well represented, and the septa are not visible. I was inclined to regard this form as only a variety of F. subrotunda, but Dr. Sterki differs from me in this, and he calls my attention to the fact that the color of the gonads in F. ebena is not crimson, as in sub- rotunda, but distinctly purple. I have been able to verify this. Al- though I had before me not more than half a dozen F. ebena, and although I found the gonads white in some, the others had them more or less (lighter or darker) purple, differing distinctly in hue from speci- mens of subrotunda, which I had at hand simultaneously. This matter, however, should be further investigated. In other respects F. ebena agrees with F. subrotunda, and also has the two types of color of the soft parts, whitish and orange. I have never seen gravid females, but Lefevre and Curtis (1910, p. 97, fig. 1) have figured the glochidium, which is identical with that of F. sub- rotunda, and has practically the same dimensions (length 0.14; height 0.15). Fusconaja kirtlandiana (Lea). Numerous specimens, chiefly from the Beaver drainage in western Pennsylvania have been investigated, among them only one gravid female with glochidia (found in the beginning of August). This agrees in every detail with F. subrotunda. The only difference is that the orange type of color of the soft parts israther infrequent, while the whitish prevails. The only gravid female was of the orange type, and had crimson gills. Glochidia identical. Length 0.13; height 0.15 mm. (see Ortmann, 19110, pl. 89, fig. 1). I am very much inclined to consider this as being only a variety of F. subrotunda. Genus CRENODONTA Schlueter. 1836. Simpson, 1900), p. 766 (as section). Shell rounded, ovate, subquadrate, or trapezoidal, with more or less elevated beaks, well developed hinge-teeth, and rather deep beak- 246 ANNALS OF THE CARNEGIE MUSEUM. cavities. Outer surface with a peculiar sculpture: heavy, oblique folds run across the disk chiefly in its posterior half (it may be that these folds are continuations of the posterior angle of the bars of the beak sculpture). Epidermis lighter or darker, brown to blackish, without distinct rays. Beak-sculpture either simply concentric, slightly angled upon the posterior ridge, and disappearing toward the disk, or continued upon the disk in a zig-zag pattern, much broken up, and irregular. The soft parts are primitive in structure. Supra-anal separated from the anal, but the mantle-connection between them very short and often absent. Inner lamina of inner gills free. All four gills are marsupial, but there is a slight differentiation in the structure of the inner and outer gills, the water-tubes of the inner gill being slightly wider than those of the outer gill. When gravid, the ovisacs expand a little more, so that their lumen becomes trans- versely enlarged, giving to the placente a compressed, leaf-like shape. Placente whitish, not very solid, and not persistent, and the glochidia are discharged in loose masses. Glochidia small, subovate, without hooks. Type C. plicata (Say).” Crenodonta approaches the following genus more than the preceding, in fact, it is very closely allied to Quadrula. The chief differential character is the sculpture of the shell. Crenodonta plicata (Say). Of this species, which is commonly called Quadrula hippopea (Lea). I have investigated numerous specimens from the shores of Lake Erie in Pennsylvania and Ohio. Gravid females were found in July; 1910, but only eggs were present in them, and no glochidia. Since there was at hand much more complete material of the follow- ing form, of which this is undoubtedly only a local race, I prefer to only give particulars of the anatomy of C. undulata, here only stating that C. plicata is absolutely identical with it in every respect. Crenodonta undulata (Barnes). A large number of specimens from the Ohio drainage in western Pennsylvania are at hand. This species is tachytictic, and the breed- 11 J have never seen it connected, although it is said to be so sometimes. 12 This species has been misunderstood hitherto. The type locality of plicata is Lake Erie, and thus the only known Crenodonta from Lake Erie should bear this name, but this is the form called hippopea by Lea. The plicata of authors (incl. Simpson) should be Cr. peruviana (Lamarck). ORTMANN: FAMILIES AND GENERA OF NAJADES. 247 ing season lastsfrom the middle of May to the middle of July. The discharge of the glochidia has been observed on July 8, 1909. The soft parts have been described by Lea (Obs., X, 1863, p. 417), but incorrectly in several particulars. They also have been described by Simpson (in Baker, 1898, p. 82). Margin of the mantle drawn together by the gill-diaphragm, thus separating the anal and branchial openings. Anal and supra-anal separated by a very short mantle-connection, which is sometimes absent (torn?). Branchial opening with strong papille, anal also with papille, which, however, are much finer, and sometimes appear only as crenu- lations. Palpi of the usual shape, their posterior margins connected only at base or up to one-third of the length. Gills broad, the inner the wider, their anterior ends as usual. Dia- phragm normal, and inner lamina of the inner gills free from the ab- dominal sac, except at the anterior end. Gills with well-developed septa and water-tubes, as usual. In the female, marsupial structure is observed in all four gills, the septa being better developed, with folded epithelium, closer together, and the water-tubes being narrower. Yet in the outer gill the water-tubes are somewhat narrower than in the inner gill, which is chiefly noticeable at the base of the gills. In the gravid female, the gills swell moder- ately, so that the ovisacs assume a lanceolate, leaf-like shape, while the edges of the gills remain sharp and do not distend. Eggs whitish, filling the ovisacs in rather poorly connected masses, although a pla- centa-like cohesion is seen. But later on this placenta-structure is lost, and the glochidia are discharged in rather loose, irregular masses. Glochidia of suboval shape, without hooks. Length 0.21; height 0.22 mm. (see Lea, Obs., VI, 1858, pl. 5, fig. 22, but not quite correct in shape; Ortmann, 1911), pl. 89, fig. 3). Color of soft parts whitish, foot, margin of mantle and gills pale brownish or yellowish. No trace of any brilliant colors (red or orange). Crenodonta perplicata (Conrad). One male, and two females, from Bayou Pierre, De Soto Parish, Louisiana, collected Aug. 6, 1910, have been received from L. S. Frier- son. One of the females proved to be gravid, and in the act of dis- charging glochidia. Three males, three females, and two young ones from Ouachita River, Arkadelphia, Arkansas, have been sent by H. E. Wheeler. 248 ANNALS OF THE CARNEGIE MUSEUM. This form is very likely the southern representative of the foregoing. It agrees with itin every respect. In one of the specimens from Louisi- ana and three from Arkansas the supra-anal was separated from the anal, in the others this separation was absent. The inner edge of the anal is finely crenulated. The posterior margins of the palpi are connected for about one-third of their length, and the inner lamina of the inner gills is free {rom the abdominal sac. The gill-structure of the females is identical with that of C. undulata. The gravid female had only a few glochidia in the outer gills, while the inner ones were yet partly charged, and both suprabranchial canals, as well as the cloacal chamber, were filled with masses of loose glo- chidia, partly sticking together, but not in the shape of placente. Glochidia like those of C. undulata. Length c.20; height 0.21 mm. Crenodonta heros (Say). According to the description and figure given by Lea (as multi- plicatus, Obs., VII, 1860, p. 222, pl. 30, fig. 105), this species without doubt belongs here. Crenodonta trapezoides (Lea). I have received, from L. S. Frierson, one male and two females from Bayou Pierre, De Soto Parish, Louisiana (collected Aug. 6, 1910), Fic. 5. Crenodonta trapezoides (Lea). Male, from Bayou Pierre, De Soto Parish, La. (Carn. Mus., No. 61, 4,586.) from A. A. Hinkley two females from Pearl River, Jackson, Hinds Co., Mississippi (collected Nov. 5, 1910), and from H. E. Wheeler a male ORTMANN: FAMILIES AND GENERA OF NAJADES. 249 anda female from Ouachita River, Arkadelphia, Clark Co., Arkansas. None of the females was gravid. The description of the soft parts given by Lea (Obs., X, 1863, p. 436) is incomplete. The anal opening is separated from the supra-anal by a moderately long connection of the margins of the mantle, which varies slightly, and is a little longer than the anal, but always much shorter than the supra-anal. In two cases this connection was absent. Branchial with well developed papilla, anal with minute papilla. Inner lamina of inner gills free, except at the anterior end. Posterior margins of palpi connected for about one-half of their length. Fic. 5a. Left gills of a sterile female of C. trapezoides, from Pearl River, Jackson, Hinds Co., Miss. (Carn. Mus., No. 61, 4,924.) Septa of the gills of the male rather distant from each other, and water-tubes wide. In the female the septa are much more crowded, and the water-tubes are narrow, chiefly so in the outer gill. In the inner gill of the female, near the base, the septa are a little more distant, but toward the edge they become more crowded by intercalation of additional ones, so that in the marginal half of this gill the water- tubes are almost as narrow as those of the outer gill. Altogether the marsupial character of the crowded septa is not so distinctly pro- nounced in the inner gill, although all four gills are built to receive eggs and to serve as marsupia, a fact, which is evidenced by the struc- 250 ANNALS OF THE CARNEGIE MUSEUM. ture of the septa and the epithelium, which is folded and wrinkled as usual in marsupial gills. Soft parts whitish. Frierson writes to me that in this species “eggs are in two gills sometimes, mostly in all four.’’ Of those sent to me (five females), the structure of the gills was alike, in every case all four gills had the marsupial structure as described above. In this species I see the nearest approach to a tendency to restrict the marsupial function to the outer gills, in so far as the water-tubes are slightly wider in the inner gills. But still the latter partake in the formation of the marsupium, and are distinctly marsupial at least in their marginal half. This species stands rather isolated also with regard to the characters of the shell, but the features of Crenodonta are clearly seen, and I think that the most closely allied form is C. heros. Genus QUADRULA Rafinesque. 1820. Simpson, 1900), p. 765 (restricted). Shell rounded, quadrate, or subrhomboidal, sometimes elongated, with rather high beaks, well developed hinge-teeth, and deep beak- cavities. Outer surface more or less sculptured, with tubercles, pustules, or ridges, but without the characteristic oblique folds of ‘Crenodonta. Epidermis lighter or darker, generally with rays, and often with beautiful color-patterns caused by the breaking up of the rays. Beak-sculpture concentric, double-looped, or zig-zag, poorly developed, or extending upon part of the disk. Soft parts primitive in structure. Supra-anal separated from the anal by a short mantle-connection, the latter sometimes absent. Inner lamina of inner gills free. All four gills marsupial (see Plate XVIII, fig. 1), but the water-tubes of the inner gills sometimes a little wider than those of the outer gills, although this difference is occasion- ally hardly noticeable. When gravid, the ovisacs expand moderately, giving a compressed, leaf-like shape to the placente. Placente (where known) whitish, not very solid, and not persistent. Glochidia, in the few cases known, small, or medium, subovate, without hooks. Type Q. metanevra (Rafinesque). The species belonging to this genus may easily be separated into three groups: 1. Pustulosa-group. Shell more or less rounded and swollen over the disk, with pustules ORTMANN: FAMILIES AND GENERA OF NAJADES. 251 (rarely smooth), which are irregularly scattered, and have no connec- tion with the beak-sculpture. Beak-sculpture poorly developed, simply concentric. QO. pustulosa, spherica, refulgens, mortont. 2. Lachrymosa-group. Shell subquadrate or subtrapezoidal, sometimes somewhat elongate. Generally profusely sculptured upon the disk with tubercles, pustules, and ridges of a more definite arrangement. This sculpture is contin- uous with the beak-sculpture, which is quite distinct, and of the double- looped type. Shell with a rather distinct, but narrow, posterior ridge and in front of this flattened, or with a broad and shallow de- pression. Epidermis rayed, rays not broken, but irregular, and with the tendency to spread over the epidermis. Q. lachrymosa, aspera, tuberculata. 3. Metanevra-group. Shell subquadrate, or subtrapezoidal, sometimes quite elongated. Surface generally sculptured with tubercles and pustules of a more or less distinct arrangement, the sculpture continuous with the beak- sculpture, which is of the double-looped or zig-zag type. Shell with a distinct, broad, and high posterior ridge, depressed in front of this, but without distinct and broad radial furrow. Epidermis rayed, rays broken up into characteristic triangular spots. Q. metanevra, sparsa, cylindrica. Quadrula pustulosa (Lea). I have investigated about half a dozen specimens from the Ohio drainage in western Pennsylvania, West Virginia, and Ohio, and (var. schoolcraftensis Lea) from Lake Erie in Ohio. Further I had ten specimens, males and females, from Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler). Both males and females were seen, but none of the latter gravid. ; The soft parts have been described by Simpson (in Baker, 1898, p. 87). Anatomy similar to that of the genera Fusconaja and Crenodonta, chiefly the latter. Anal and supra-anal separated by a short mantle- connection. I never found the latter absent. Branchial opening with papilla, anal crenulated. Palpi of the usual shape, their poste- rior margins connected for about one-third of their length. Gills short and broad (according to shape of shell), the inner gill 252 ANNALS OF THE CARNEGIE MUSEUM. the wider. Anterior attachment of gillsas usual. Diaphragm normal. Inner lamina of inner gills free from abdominal sac except at anterior end. Gills with well-developed septa, which are rather distant in the male. In the female, all four gills are marsupial, the septa being close together, and the water-tubes narrow. In the inner gill the septa are slightly less crowded near the base, but they have the characteristic marsupial structure (see Ortmann, 191Ia, pl. 7, fig. 1). The glochidia are figured by Lefevre and Curtis (1910, p. 97, fig. F). Length 0.23; height 0.32, which is unusually large for this group of genera. The color of the soft parts is grayish, or yellowish white. Quadrula spherica (Lea). Three sterile females from Pearl River, Jackson, Hinds Co., Missis- sippi, are at hand, collected on Nov. 5, 1910, by A. A. Hinkley. Structure essentially as in Q. pustulosa. Anal opening with fine crenulations, almost smooth. In all three specimens all four gills possess the marsupial structure, and the water-tubes of the inner gills are also not quite so narrow as those of the outer gills, chiefly near the base. By its shell this species is very closely allied to the foregoing, and the soft parts are practically identical. JI hardly think they are speci- fically distinct, and among the specimens of pustulosa from Ouachita River, mentioned above, there are intergrades between the two forms, Quadrula refulgens (Lea). One male, collected together with Q. spherica, is before me. Since there is only a male, the characteristic Quadrula-structure cannot be made out. But I have no doubt that this is a Quadrula on account of its close affinity to Q. spherica. In fact all the details, both of the shell and the soft parts, are identical with the Jatter, except that the shell is more compressed (lenticular) in Q. refulgens. My specimen is more rounded in outline than the original figure of Lea, and thus more nearly approaches Q. spherica in this character. I should not be astonished, if refulgens should turn out to be a mere “form” of spherica. ORTMANN: FAMILIES AND GENERA OF NAJADES. 253 Quadrula mortoni (Conrad). Three males and two females, one of the latter gravid, from Bayou Pierre, De Soto Parish, Louisiana, collected by L. S. Frierson, Aug. 6, 1910. Agreeing in every detail with pustulosa and spherica, to which it is allied. The inner edge of the anal opening is almost smooth. In the gravid female, eggs and glochidia were present, the latter of the usual shape, of medium size, subovate, without hooks. The glochidia are quite young and their shape is not very distinctly seen. They were of whitish color, and distributed in an irregular way in certain ovisacs both of the outer and inner gills, many ovisacs being empty. No exact measurements of the glochidia can be given. The date for the breeding season should be noted. Quadrula lachrymosa (Lea). One male and four females from the Wakarusa River, Lawrence, Douglas Co., Kansas, received from R. L. Moodie, and one female collected by myself in the Ohio River, at St. Marys, Pleasants Co., West Virginia. No gravid females have been seen. Soft parts described by Simpson (cf. Baker, 1898, p. 84). Similar to the preceding species. Inner edge of the anal opening irregularly and indistinctly crenulated, almost smooth. Posterior margins of palpi connected for over one-half, almost two-thirds, of their length. All four gills are marsupial, septa of the inner ones slightly less crowded than those of the outer ones (see Plate XVIII, fig. 1). Soft parts whitish. Quadrula aspera (Lea). Three males, one female (all small, or of medium size) from Bayou Pierre, De Soto Parish, Louisiana, collected by L. S. Frierson, and one very large female from Pearl River, Jackson, Hinds Co., Mississippi, collected by A. A. Hinkley. An incomplete description is given by Lea (Obs., X, 1863, p. 437). Agreeing in every respect with Q. lachrymosa, to which it is closely allied. Even the minor details (anal opening, palpi, etc.) are abso- lutely identical. No gravid females have been seen. 254 ANNALS OF THE CARNEGIE MUSEUM. Quadrula tuberculata (Barnes). Fifteen specimens have been investigated, collected by myself in the Ohio drainage in western Pennsylvania; nine more have been re- ceived from H. E. Wheeler from the Tennessee drainage in northern Alabama, and the Ouachita River in Arkansas. Females are among them, but not in the gravid condition. Simpson has created for this species the genus Tritogonia, which he removed far from Quadrula. The shape of the shell is indeed somewhat strange at the first glance, but it is possible, without much difficulty, to correlate shape and sculpture with that of such species as Jachrymosa, aspera, and chiefly with certain southern forms, which probably also belong here (forshei Lea, speciosa Lea, apiculata Say). In the structure of the soft parts, this species is essentially a Quad- rula. The anal opening is separated from the supra-anal by a rather short mantle-connection; the latter was found absent in one case only (out of twenty-four). Branchial with well developed papille, anal with fine, but distinct crenulations, which sometimes resemble fine papillae. Inner lamina of inner gills free from abdominal sac, except at its anterior end. Posterior margins of palpi connected for one-half, or even more, of their length. Gills rather long, but also rather wide; their anterior attachment as usual. Septa well developed, rather distant from each other in the male. In the female they are more crowded in all four gills, and the water-tubes are narrow, but there is a slight difference between the inner and outer gill, the water-tubes of the former being slightly wider near the base of the gills. In the marginal portion there is hardly any difference in the water-tubes of the two gills (see Ortmann, 19110, pl. 86, fig. 4). In all four gills the septa are distinctly marsupial in structure: they are heavy, and have a folded epithelium. No gravid females have been seen by the writer, and the glochidia are still unknown. The color of the soft parts is grayish or yellowish (or brownish) white. Simpson (1900), p. 608) says of his genus Tritogonia: ‘‘in the female there is a thickened flap of the mantle which fills the circular posterior expansion of the shell, and which has a small flap inside.’”’ I have never seen anything answering to this phrase in my specimens. The chief expansion of the shell is at the anal opening, and the margin of this opening corresponds to it, and thus the anal is larger in the female, ORTMANN: FAMILIES AND GENERA OF NAJADES. 255 than in the male. The ‘‘inside flap’’ can only be the inner edge of the mantle, which is present, however, in the male also. The true position of ‘this species was first indicated by Sterki (1907, p. 48). Quadrula metanevra (Rafinesque). Thirteen specimens of either sex, one a gravid female, were exam- ined in the laboratory, all from the Allegheny and Ohio Rivers in western Pennsylvania; ‘additional specimens were examined in the Fic. 6. Quadrula metanevra (Rafinesque). Male, from Allegheny River, Kelly, Armstrong Co., Pa. (Carn. Mus., No. 61, 4,549.) Coll. May 20, 1910. field, in the Ohio River in West Virginia and Ohio, and two (male and female) were received from the Ouachita River in Arkansas (H. E. Wheeler). The gravid female was found on June 22, 1909, and had only eggs, Margins of the mantle con- nected so as to separate anal and supra-anal openings; this connection is short, but in no case was found to be absent. Supra-anal very large. Bran- chial with well developed pa- Fic. 6a. Left gills of a sterile female, from same locality. pille, anal practically smooth, or only with mere traces of ir- regular crenulations. Palpi with the posterior margins connected for about one-fourth to one-third of their length. Gills short and wide, their anterior attachment as usual. Dia- 256 ANNALS OF THE CARNEGIE MUSEUM. phragm normal. Inner lamina of inner gills free from abdominal sac, except at anterior end. Septa and water-tubes well developed. In the female all four gills are marsupial, and possess the typical structure. In the basal portion of the inner the water-tubes are somewhat wider, but there is hardly any difference in their width in the marginal part of the two gills, since the water-tubes of the inner gills become narrower by inter- calation of additional ones. In the gravid female the gills swell moderately, but their edges remain sharp. The eggs form only poorly developed placenta in the ovisacs, and the shape of the latter is com- pressed and lanceolate (leaf-like). The eggs are whitish. I have not seen glochidia, but according to Lefevre and Curtis (1910, p. 97, fig. E) they are normal in shape and size. Length 0.18; height 0.19 mm. Color of soft parts whitish. As usual, the edge of the mantle, chiefly along the posterior part, is more or less blackish or brownish. Gills paler or darker grayish or brownish white. Foot brownish white. The posterior part of the abdominal sac is often suffused with black. Quadrula sparsa (Lea). One male and one sterile female, from the Cumberland River in Cumberland and Pulaski Counties, Kentucky, at hand, received from B. Walker. . Identical in every detail with Q. metanevra, to which it is also allied by the shell. The agreement extends so far, that minor details are also identical, as the smooth edge of the anal, the shape of the palpi, and the black pigment of the posterior part of the abdominal sac. In the male supra-anal and anal were not separated, but this region was somewhat injured, so that the mantle-connection may have been torn. Charged marsupia and glochidia unknown. Quadrula cylindrica (Say). Nine specimens (males and females) from the Ohio drainage of western Pennsylvania have been examined in the laboratory, and several more in the field, taken from the Ohio River in western Penn- sylvania and Ohio. Two males were received from H. E. Wheeler, from the Ouachita in Arkansas. ' ORTMANN: FAMILIES AND GENERA OF NAJADES. 257 Although the shape of the shell is very unique in this species, it clearly belongs to the metanevra-group, as has been recognized already by Simpson, and the soft parts bear out this affinity, since they are identical in all essential characters. Of course, according to the shape of the shell, the gills are very long and narrow. The mantle connection between anal and supra-anal was always found present. The inner edge of the anal is practically smooth. The posterior margins of the palpi are united for about one-half of their length. All four gills have a marsupial structure in the female, and the inner gill has the water-tubes a little wider in the basal part than the outer gill. No gravid specimens have been found. The color of the soft parts of this species is very remarkable. The general ground color is yellowish orange, with black markings. The abdominal sac is whitish, with blackish gray markings, chiefly pos- teriorly. The foot is grayish at the extremity, shading into black, the black ending in a sharp horizontal line, which is followed by grayish orange. The palpi are yellowish, with a gray edge, the gills are grayish brown, the mantle transparent gray, shading to grayish yellow on the margin, with a blackish brown edge, which becomes deep black and very wide posteriorly at the siphons. The adductors are pale yellowish to orange. The color varies in different specimens in so far that in some the ground-color is paler yellow, while in others it is of a more intense yellow, inclining to orange. Breeding season and glochidia unknown. Genus ROTUNDARIA Rafinesque. 1820. Simpson, 1900), p. 794 (as subgenus). Shell rounded or quadrate, with elevated beaks, very deep beak- cavities, and well developed hinge-teeth. Outer surface sculptured with tubercles and nodules. Epidermis brown, without rays. Beak- sculpture consisting of numerous rather close bars, the first few con- centric, those following developing a strong angular loop on the posterior ridge, and an anterior loop, which soon breaks up into an irregular zig-zag pattern of more or less isolated tubercles. This sculpture extends somewhat upon the disk and mingles with the first tubercles of the disk. Nacre more or less violet, which may be a specific character. Soft parts primitive in structure. The anal is never closed above, 258 ANNALS OF THE CARNEGIE MUSEUM. and no supra-anal is present. Inner lamina of inner gills free. Of the gills, only the outer ones are marsupial in the female, and the water- tubes in them are much closer than in the non-marsupial gills. Glo- chidia unknown. Type R. tuberculata (Rafinesque). This genus is more primitive than any of the foregoing because of the absence of a supra-anal opening. In the structure of the mar- supium it is a little more advanced, and is related to the following genera. In the characters of the shell it is rather peculiar, but re- minds somewhat of certain types of Quadrula. We can do justice to these conflicting characteristics only by recognizing this as a valid genus. Rotundaria tuberculata (Rafinesque). Three males and three females from the Ohio drainage in western Pennsylvania, and one female from the Ohio River at St. Marys, Pleasants Co., West Virginia, all collected by myself, have]been investigated. o=mp Fic. 7. Rotundaria tuberculata (Rafinesque). Sterile female, from Allegheny River, Kelly, Armstrong Co., Pa. (Carn. Mus., No. 61, 4,562.) Coll. July 25, 1910. The soft parts have been described by Simpson (in Baker, 1898, p. 86), but very poorly. Although there are only a few specimens at my disposal, the char- acters given for the genus are uniformly found in all of them. The most marked character is the absence of a supra-anal. Since Simpson ORTMANN: FAMILIES AND GENERA'OF NAJADES. 259 also mentions this character, I think we may take it as settled that this form does not possess a supra-anal. As regards the marsupium, I can only say that in all four females at hand only the outer gills show the marsupial structure. In conclusion there is nothing remarkable in the structure of the soft parts. The branchial opening is unusually large, and has papille on the inner edge, while the large anal has a practically smooth inner edge. The palpi are normal, and their posterior margins are united for one-third to one-half of their length. The gills, conforming to the shape of the shell, are rather short and wide, and their anterior ends are normal, the diaphragm is complete, and the inner lamina of the inner gills is free from the abdominal sac, except at the anterior end. The gills have the usual structure, and the female marsupial structure is found only in the outer gills, where the septa are much more crowded, ‘and the water-tubes much narrower, than in the non-marsupial gills. A gravid female has never been found, and consequently the glo- chidia remain unknown. The color of the soft parts is grayish or brownish white, with ex- ception of the margin of the mantle, which is brownish, and becomes deep black in the region of the branchial and anal openings. Genus PLETHOBASUS Simpson. (1900.) Simpson, 1900), p. 764 (as section). Shell rounded, oval], or slightly elongate, with moderately elevated beaks, and moderately deep beak-cavities, and well developed hinge- teeth. Outer surface sculptured with nodules or tubercles, which often are transversely elongated. Epidermis yellowish to brown, without distinct rays. Beak-sculpture rudimentary, consisting (as far as known) of a few concentric ridges, which do not extend upon the disk. Nacre whitish or pinkish. Soft parts primitive in structure. Anal separated from the supra- anal by a short mantle-connection. Inner lamina of inner gills free. Only the outer gills are marsupial in the female, in other respects the gills have the usual structure. When gravid, the outer gills swell moderately, and their edges do not distend. Placente lanceolate (leaf-like) and compressed; they are rather persistent, and are dis- charged whole. Glochidia small, semioval, without hooks. Type P. esopus (Green). 260 ANNALS OF THE CARNEGIE MUSEUM. Possibly the color of the soft parts and of the eggs (placentz) is also characteristic. (See below.) This genus greatly resembles in shell-structure the genus Quadrula, chiefly the pustulosa-group and the genus Rotundaria. But in the marsupium it is more advanced than the former, and in the color of the soft parts and the eggs it suggests affinity with Fusconaja. It surely is a connecting link between the more primitive Unionine and those of the type of the genus Pleurobema. Simpson has recognized the peculiar character of P. @sopus, and I think his section Pletho- basus is entitled to generic rank. Plethobasus zsopus (Green). Some fifty specimens have been examined, chiefly from the Allegheny River in Armstrong Co., Pennsylvania. Among them were gravid females, and the latter were found exclusively in the month of July. : The species is typically tachytictic. OoO=mp Fic. 8. Plethobasus @sopus (Green). Sterile female (just discharged), from Allegheny River, Kelly, Armstrong Co., Pa. (Carn. Mus., No. 61, 4,598.) Coll. July 25, 1910. Anal and supra-anal separated by a very short mantle-connection, which was found missing in a small number of cases. Branchial opening with strong papilla, anal with very fine papilla. Palpi of usual shape, their posterior margins connected for about one-third of their length. Gills long and broad, the inner the wider. Anterior attachment as usual. Diaphragm normal. Inner lamina of inner gills free. Septa ORTMANN: FAMILIES AND GENERA OF NAJADES. 261 and water-tubes well developed, septa rather distant in the male. In the female, only the outer gill is marsupial, with crowded septa, while in the inner gill the septa are like those of the male. When gravid, the water-tubes (ovisacs) of the outer gill swell mod- erately, and assume a lanceolate shape, and consequently the placente have this shape. The edge of this gill does not distend. The eggs and placente are pink or red, and the placente preserve their shape, and are discharged whole. This discharge has been actually observed in two cases (July 25, 1910). Glochidia of the usual shape, rather small, semioval, without hooks.!8 Color of soft parts very characteristic. All specimens seen had a peculiar, pale orange ground-color. The foot, margins of the mantle, and adductors were darker, often deep orange. The abdominal sac is whitish, the palpi and gills pale grayish brown, the gills with more or less of the orange hue. The color may be more or less intense, but some shade of orange always prevails. When gravid, the red color of the placente contained in the gills blends with the orange of the latter, producing a very peculiar lilac tint. Plethobasus cooperianus (Lea). Only two specimens with soft parts have been observed, the one from the Ohio River in Beaver Co., Pennsylvania, the other from the Ohio at Parkersburg, Wood Co., West Virginia. Both proved to be females. The anatomy of this species is practically identical with that of P. esopus agreeing in all particulars, chiefly also in the peculiar color of the soft parts. This color is so characteristic that it alone suggested to me the relationship of cooperianus and @sopus, which was confirmed by the subsequent anatomical investigation. No gravid females have been observed, and nothing is known about the color of the placentae and the shape of the glochidia. Genus PLEUROBEMA Rafinesque. 1820. Simpson, 1900), p. 745 (amended). Shell rounded, subquadrate, oval, or somewhat elongate, but then oblique, with the beaks placed anteriorly. Beaks generally somewhat 13 The glochidia were observed only in one case, in one of the discharging females, and apparently were discharged prematurely (a phenomenon observed by others). They were young and incompletely formed (soft), but their shape could be made out. The size was about 0.18 mm., but it was impossible to make exact measurements. 262 ANNALS OF THE CARNEGIE MUSEUM. prominent, with moderately deep beak-cavities; hinge-teeth well developed. Outer surface without sculpture. Epidermis generally lighter or darker brownish, sometimes yellowish, with more or less distinct rays, which may be arranged in blotches, or may be fine lines, or entirely absent. Beak-sculpture obscure, consisting of a few concentric ridges, not extending upon the disk. Nacre generally whitish, or red. Soft parts practically identical with those of Plethobasus, except in color, which is generally paler, sometimes yellowish, even with a suggestion of pale orange (in P. clava), but in most cases without this. The eggs are, where known, white, and the placente are distinct, and seem to be persistent. Glochidia small, suboval, without hooks. Type P. clava (Lamarck). It isan unhappy coincidence, that the type species does not represent the normal condition of the genus, but a rather extreme form of it. In general, we may say that Pleurobema resembles Quadrula in the soft parts, and Fusconaja in the shape of the shell, with the exception that it has reached the advanced stage of having only the outer gills marsupial. It differs from the following genera (Elliptio and Unio- merus) only by the characters of the shell, chief among which is the general outline, which is of the short, rounded, quadrate, or oblique type, while Elliptio and Uniomerus have the elongate and straight type of shell. Simpson (1900), p. 760, footnote 2) had difficulty in defining this genus, and there are indeed species which are to a degree intermediate between Pleurobema and Elliptio. PP. clava is a peculiar type. Since I have not had any opportunity to investigate these doubtful forms, I have made the present arrangement of the genera to suit the material at hand, but I shall not be astonished if further studies will necessitate changes. Pleurobema riddelli (Lea). One young male, and one larger female (sterile) are before me, from Pearl River, Jackson, Hinds Co., Mississippi, collected by A. A. Hinkley. I was a little uncertain about the identification of this species, but since Mr. Walker, to whom I sent these specimens, also thinks that they are riddell, | believe the identification is right. 14 The species of Pleurobema described by Rafinesque are unrecognizable, but L. Agassiz (1852) has made this the type. ORTMANN: FAMILIES AND GENERA OF NAJADES. 263 Structure of the soft parts like those of the following species, which see. Color of soft parts also of similar type. Mantle-connection separating anal and supra-anal very short. Anal with fine crenula- tions, branchial with papillae. Posterior margin of palpi connected for about one-half of their length. Gills of the usual structure, in the female only the outer gills are marsupial (see Plate XVIII, fig. 2). In the soft parts there are no characters which assign this species to a definite genus, except that they indicate, that it is not Fusconaja and not Quadrula. It has been placed by Simpson in the latter genus. In the shape of the shell it most resembles the coccineum-obliquum- group of Pleurobema, from which it is distinguished by the rather distinct posterior ridge. I think, P.riddelli comes in here, but stands rather isolated by itself. The dark color of the epidermis is also un- usual in this genus. Pleurobema coccineum (Conrad). Numerous specimens have been examined from the smaller creeks of the Ohio and Lake Erie drainage in western Pennsylvania, among them some fifty gravid females. The species is tachytictic, and the breeding season lasts from the end of May to the end of July. The soft parts have been described by Simpson (in Baker, 1898, p- 79), but the account given of the marsupium is wrong. Margin of the mantle, branchial, anal, and supra-anal openings of the usual conformation, as also the gills and palpi. Branchial with papilla, anal with distinct, almost papille-like crenulations. The mantle-connection between the anal and supra-anal was found absent in some cases. Posterior margins of palpi connected for one-half, or even more, of their length. Only the outer gills are marsupial.’ When gravid, the outer gills swell only moderately, the edges remaining sharp. The ovisacs are compressed and lanceolate and the placenta, which are distinctly developed, have the same leaf-like shape; they are always white, and are permanent, being discharged whole. This discharge has been observed several times. Glochidia rather small, subovate, without 15 Lefevre and Curtis (1910, p. 83) suggest that the fact that I found only the outer gills charged may be due toa partial discharge of the marsupia in consequence of beginning suffocation. This supposition is untenable. I observed many speci- mens in the field. Moreover, even in females, which have the marsupium not charged, it is easily seen that only the outer gills have marsupial structure. 264 ANNALS OF THE CARNEGIE MUSEUM. hooks (see Ortmann, 19110, pl. 89, fig. 4). Length and height about the same: 0.15 mm. Color of soft parts grayish or yellowish white. Among the numerous specimens investigated not one has been found which showed any traces of orange color. Pleurobema obliquum (Lamarck). A large number of specimens from the Ohio and Allegheny in Pennsylvania, and the Ohio in West Virginia and Ohio have been investigated. Gravid females have been found only a few times in June. Structure of soft parts absolutely identical with that of P. coccineum, but glochidia have not been observed. I do not think that this form is specifically distinct from P. coc- cineum. It is the form of the large rivers, which is represented in the headwaters and smaller streams by P. coccineum. In the Allegheny River in Armstrong Co., Pennsylvania, these two forms are connected by all kinds of intergrades. Pleurobema pyramidatum (Lea). Not more than a dozen specimens of typical pyramidatum have been seen, found always associated with the foregoing form. Females were among them, but none gravid. I also received one male and three females of this form from Arkadelphia, Arkansas, collected by H. E. Wheeler. This is merely an extreme variety of P. obliquum, connected with it by frequent transitional forms, and consequently the anatomy is absolutely identical. Pleurobema clava (Lamarck). About twenty-five specimens, among them gravid females, have come under observation. They are all from the Ohio drainage in western Pennsylvania. This species is gravid in June and July. The soft parts have been described by Lea (Obs., X, 1863, p. 441), but only those of the male. Anatomy like that of the other species of Plewrobema. It should be mentioned that the mantle-connection between the anal and supra-anal is rather short, and was always found present. The anal is rather distinctly, but finely, papillose. Posterior margins of palpi connected for only a short distance. ORTMANN: FAMILIES AND GENERA OF NAJADES. 265 The outer gills alone are marsupial, and the placente are rather distinct. Glochidia (see Ortmann, 19110, pl. 89, fig. 5) of small size, subovate, without hooks. Their length and height is about the same, 0.16 mm. Color of so{t parts whitish, with foot and gills grayish, and the margin of the mantle black posteriorly. In other specimens the foot Wy mp me U7) fe) Fic. 9. Pleurobena clava (Lamarck). Male, from Sandy Creek, Sandylake, Mercer Co., Pa. (Carn. Mus., No. 61, 4,589.) Coll. June 27, 1910. Fic. 9a. Left gills of sterile female, from Shenango River, Pulaski, Lawrence Cot;, Pa. “(Gam Muss No: 6m, 4,501.)). ‘Coll. Oct. 4; roro: is pale orange, as are also the margins of the mantle and adductors. The gills are grayish brown. There are all intergrades between these extremes. The placenta are white, cream-color, or pale orange. Pleurobema decisum (Lea). According to the similarity of the shell, and the notes on the anatomy furnished by Lea (Obs., X, 1863, p. 405), this species belongs here. Genus ELiipTio Rafinesque. I819. Simpson, 1900), p. 700 (as section). Shell more or less elongated, with straight longitudinal axis, not oblique. Beaks not very near the anterior end, not very prominent, with shallow beak-cavities; hinge-teeth well developed. Outer surface without sculpture.!® Epidermis generally rather dark, brown to black, without rays, or with indistinct rays, chiefly so when young. Rays straight, simple, and fine. Beak-sculpture tending to become If U. spinosus belongs here, the diagnosis should be modified in this par- ticular. 266 ANNALS OF THE CARNEGIE MUSEUM. obsolete; when present, consisting of few fine concentric ridges, to which may be added, toward the disk, a small number of slightly heavier bars, with a posterior angle upon the posterior ridge of the shell; these bars run about parallel to the growth lines. Sometimes the bars are slightly sinuate in front of the posterior angle, but they never are distinctly of the double-looped type. Nacre from white through all shades of pink and red to deep purple and violet, with the dark shades prevailing. Soft parts practically identical with those of Plethobasus and Pleuro- bema, with only the outer gills marsupial. Mantle-connection between anal and supra-anal short, or somewhat longer. Inner lamina of inner gills free. Color of the soft parts whitish, often greatly suffused (chiefly the gills and mantle) with black. Eggs, where known, whitish, placente rather distinct. Glochidia small, subovate, without hooks (see Plate XIX, fig. 1). It may be mentioned as an additional character, that in all these forms the anal has rather distinct papille. Type: E. crassidens (Lamarck.)" We may regard Eiliptio as a special branch of Pleurobema, distin- guished from the latter only by the characters of the shell. It prob- ably is not descended directly from a Quadrula- or Fusconaja-like type with four marsupial gills, but it has gone through the intermediate Pleurobema-stage first. The species of Elliptio easily fall into several groups. In E. crassidens and beadleianus, the typical shape of the shell is not so distinctly developed: it is not greatly elongated, and approaches yet somewhat the subquadrate or subtrapezoidal type of certain Fusconaja- and Pleurobema-species. The beak sculpture in these forms is rather obscure. Next to this stands the group of E. complanatus (including productus and jayensis), where the typical characters of the genus are fully de- veloped. A third type is furnished by E. gibbosus, and a fourth by E. popet. Elliptio crassidens (Lamarck). Numerous specimens both males and females have been examined, all from the Ohio and Allegheny Rivers in western Pennsylvania. 1 Unio (Elliptio) nigra Rafinesque, 1820, is Rafinesque’s type (first species), and this is undoubtedly a synonym of U. crassidens Lamarck, 1819. The large, heavy shell of the Ohio with red nacre cannot be anything else. ORTMANN: FAMILIES AND GENERA OF NAJADES. 267 There were only three gravid females among them, which were found on June 22, 1909. Anatomy normal: margin of the mantle, siphons, gill-structure, and marsupium typically Unionine. In a few cases the rather short Fic. 10. Elliptio crassidens (Lamarck). Male, from Allegheny River, Kelly, Armstrong Co., Pa. (Carn. Mus., No. 61, 3,777.) Coll. July 3, 1908. Fic. 10a. Left gills of a sterile female, from same locality. (Carn. Mus., No. 61, 3,055.) Coll. Sept. 5, 1907. mantle-connection between the anal and supra-anal was found to be absent. Anal with small, but quite distinct, papilla. Palpi with the posterior margins connected for a short distance. 268 ANNALS OF THE CARNEGIE MUSEUM. Marsupium formed by the outer gills; when gravid moderately swollen, with rather well-developed, leaf-like placente. Glochidia (see Ortmann, 19110, pl. 89, fig. 6) small, suboval, without hooks. Length 0.13; height 0.15 mm. The color of the abdominal sac is whitish, the foot pale gray or brownish gray, the mantle pale liver- brown, whitish toward the margins, edge brown, black posteriorly. Gills gray or dirty brown. Adductors whitish, palpi grayish. Elliptio beadleianus (Lea). Two males and two females, from Pearl River, Jackson, Hinds Co., Mississippi, have been received from A. A. Hinkley. This species, which has been placed by Simpson (1900), p. 786) in the genus Quadrula, is not a Quadrula, because only the outer gills have marsupial structure. In other respects its anatomy is indis- tinguishable from that of other forms belonging in the genera of the type of Pleurobema, etc. The shape of the gravid marsupium, of the placenta, and of the glochidia is unknown. The supra-anal opening is well separated from the anal, but the separating mantle-connection is short. The inner edge of the anal has fine, but distinct, papilla, that of the branchial has larger papille. The posterior margins of the palpi are connected for one-third or one- half of their length (this is the most prominent difference from the allied forms). The inner lamina of the inner gills is free, as usual. Although the structure of the gills unquestionably removes this species from Quadrula and Fusconaja, it is hard to assign it a place in the other genera. We must rely entirely upon the shell, and this is rather an indifferent criterion. However, I think the shape of the shell is more like that of crassidens than that of any other form. It is some- what more elongate than the Fusconaja-Pleurobema-type, straight, with the beaks not much anterior, with a dark epidermis, and with a tendency to develop red nacre, characters which are all found in E. crassidens. The posterior ridge is also present in both species. I consider E. beadleianus a peculiar type, standing nearest to E. crassidens. Probably other species go with it, as for instance chicka- sawhensis Lea and askewi Marsh (of the latter two, Frierson writes to me that they are ‘‘next to inseparable’’). All these differ from crassidens in being smaller, possessing more regularly swollen lateral faces of the disk, lacking corrugations on the posterior slope, and having a lighter nacre. Also the whole shell and the hinge are less massive in structure. Their beak-sculpture is unknown. ORTMANN: FAMILIES AND GENERA OF NAJADES., 269 Color of soft parts of FE. beadleianus whitish; foot grayish; gills and palpi grayish-brown, as is also the mantle, except the margin, which is whitish, with the edge blackish posteriorly. Elliptio (?) spinosus (Lea). The anatomy has been described by Lea (Obs., X, 1863, p. 413). It is similar to that o! E. crassidens, but Lea mentions some peculiar- ities in the ovisacs, which I do not understand. The shell is of the crassidens-type, but its spines are unique and would possibly justify the erection of a separate genus: Canthyria Swainson, 1840. Elliptio complanatus (Dillwyn). Numerous specimens of this species, males, sterile and gravid females, have been investigated from the Delaware, Susquehanna, and Potomac drainages of eastern Pennsylvania and Maryland. The species is tachytictic, and the breeding season begins at the end of April, and lasts to the middle of June, possibly a little longer. The soft parts have been described by Lea (Obs., X, 1863, p. AID ye In the shell, this is a typical Elliptio, with the typical shape of this genus, and its typical beak-sculpture. In the soft parts, we find the edges of the mantle forming the usual openings. Anal and supra-anal are separated by a moderate mantle-connection, shorter than the anal, which has never been found missing. The branchial has large papille, while the anal has much finer ones. Posterior margins of palpi con- nected for a short distance. Gills corresponding to the shape of the shell, rather long and moderately wide, the inner rather wider. Diaphragm of the usual shape, inner lamina of inner gill free, except at anterior end. Septa and water-tubes well developed, the latter wide in the male. In the female only the outer gills are marsupial, and their septa are much crowded. When gravid, this gill swells only moderately, the edge remaining sharp, and the ovisacs assume a leaf-like shape, as also do the placente (figured by Lillie, 1895, pl. 1, fig. 1), which are not very solid when glochidia are present. The latter are always white, rather small, subovate, without hooks. They are longer than high. Length 0.20, height 0.19 mm. (see Plate XIX, fig. 1). Color of soft parts of the grayish white type, foot darker, also gills, the latter often suffused with black posteriorly. 270 ANNALS OF THE CARNEGIE MUSEUM. Elliptio jayensis (Lea). Five males and twelve females (all sterile) from Lake Monroe, Sanford, Orange Co., Florida, collected by O. T. Cruikshank, in April, 1907. The soft parts are of the usual structure, and agree in every particular with those of E. complanatus. Anal and supra-anal separated by a mantle-connection, which is slightly longer than in the more primitive forms, but still considerably shorter than the anal. Anal with rather well developed papillz, which are almost as large as the papillz of the _ branchial opening. Posterior margins of palpi connected at base only. Inner Jamina of inner gill free, except at anterior end. Only the outer gills are marsupial. The beak-sculpture of this species is nct quite of the normal Elliptio- type, in that in the case of the later bars a sinuation is seen in front of the posterior angle. This sinuation is variable, and never assumes the shape of a distinct, reéntering angle, and thus the beak-sculpture cannot be called double-looped. There are over one hundred shells (without the soft parts) at hand, many of which might as well be called E. buckleyi (Lea). All these shells undoubtedly are the same species, and were collected together. The old ones most resemble buckleyi (see: Simpson, 1892, pl. 58, figs. 6 and 7, and pl. 50, fig. 1), while those of medium size and the young ones are jayensis (Simpson, zbid., pl. 61, fig. 4). It is remarkable that there are no young buckley in the lot, while there are dozens of jayensis. JT am very much inclined to regard buckleyi as being only the older adult form of jayensis. Elliptio productus (Conrad). Nine specimens have been investigated, males and sterile females, from the Potomac drainage in southern Pennsylvania and Maryland. The soft parts agree in all essential respects with those of E. com- planatus. Conforming to the shape of the shell, the gills are extremely long and narrow. Branchial, anal, and supra-anal as in complanatus; anal and supra-anal separation slightly longer than usual; anal with the same distinct papilla. Structure of palpi and gills the same. No gravid females have been found. Color of soft parts like that of complanatus. Beak sculpture practically identical, and there is no doubt that this species is an offshoot of the complanatus-stock. ORTMANN: FAMILIES AND GENERA OF NAJADES. 271 Elliptio gibbosus (Barnes). Numerous specimens from the Ohio and Lake Erie drainage in western Pennsylvania have been seen, and two males and two females from Arkansas. Gravid females were found in the months of May, June, and July, and in one case as late as August 13. This isa typical tachytictic form. The soft parts have been discussed by Lea (Obs., X, 1863, p. 417) and Simpson (in Baker, 1898, p. 70). The beak-sculpture in this species is also similar to that of E. com- planatus, but it is slightly heavier, though less distinct; that is to say: the ridges are thicker, but less well defined. Soft parts essentially identical with the other species described in this genus. It, however, should be mentioned that in a very few cases the mantle-connection between the anal and the supra-anal was found missing. The anal has distinct papille. When gravid the marsupium swells moderately, but the edge re- mains sharp. The ovisacs are leaf-shaped, and the placente are moderately well-developed, but when the glochidia are formed, they seem to be less distinct. Color of eggs and glochidia always white. Glochidia (see Lea, Obs., XIII, 1874, pl. 21, fig. 10; and Ortmann, 19110, pl. 89, fig. 7) rather small, suboval in shape, without hooks. Length 0.20, height 0.22 mm. Color of soft parts grayish white. Foot, gills, and mantle gray, edge of the latter black posteriorly. Marsupium cream-white. Elliptio popei (Lea). Two gravid females, from Valles River, Mexico, collected by A. A. Hinkley in December and January, 1906-1907," were received from L. S. Frierson. The beak-sculpture is somewhat different from that which is typical of this genus. In the two specimens before me, it is poorly developed, although the beaks are well preserved. It consists of two to three fine concentric ridges, which are somewhat interrupted in the middle, giving a faint appearance of double loops. But it is not by any means double-looped, since no reéntering angles are present. The soft parts are typical. The anal and supra-anal are separated by a mantle-connection, which is shorter than the anal. Anal with well-developed papilla; papilla of branchial larger than those of anal. 18 See Hinkley, 1907, pp. 68 and 79. Die ANNALS OF THE CARNEGIE MUSEUM. Posterior margins of palpi connected for a short distance. Inner lamina of inner gills free, except anteriorly. Both of my specimens are gravid, but have only eggs. Marsupium formed by the outer gills, only moderately swollen, with sharp edge. Placenta moderately well developed. Hinkley collected these specimens in December and January. Here we would have a so-called ‘‘summer breeder,’’ which breeds in mid- winter. But we know now, that not the season of the year, but the shortness of the breeding season is important, and according to all analogies, E. popei should be a form with short breeding season. Genus UNIOMERUS Conrad. (1853.) Conrad, 1853, p. 268.—Simpson, 1900), p. 739 (as section). Shell moderately elongated, with straight longitudinal axis, not oblique, and beaks not very near the anterior end. Beaks not very prominent, beak-cavities shallow, hinge-teeth well developed. Outer surface without. sculpture. Epidermis light yellowish to brown, often with dark concentric bands, without rays. Beak-sculpture rather distinct, concentric, bars rather numerous, not angled behind, but curved up toward the posterior side of the beaks, and not parallel to the growth lines. Nacre whitish or grayish, not inclining to purple orred. Soft parts practically identical with those of Elliptio. Gravid females are unknown, but in sterile females only the outer gills are marsupial in structure. The anal has, in the type species, only crenu- lations, and the mantle-connection between anal and supra-anal is rather long. Type U. tetralasmus (Say).¥ This genus stands very close to Elliptio, and, like this, may be regarded as descended from Pleurobema. Uniomerus tetralasmus (Say). One male and two females (sterile) from Bayou Pierre, De Soto Parish, Louisiana, have been received from L. S. Frierson. The soft parts do not offer anything remarkable, when compared with those of Pleurobema and Elliptio. The supra-anal is separated from the anal by arather long mantle-connection, the latter, however, is 19 The first species given by Conrad is declivis, which, according to Simpson, together with six of the other so-called species named, are synonyms, or varieties, of tetralasmus. ORTMANN: FAMILIES AND GENERA OF NAJADES. ihe shorter than the supra-anal, but distinctly longer than the anal. Inner edge of anal with fine crenulations, that of branchial with papille. Inner lamina of inner gills free, except anteriorly. Posterior margins of palpi connected for about one-fourth of their length. Marsupium formed by the outer gills, and of the usual structure. Color of soft parts whitish. Genus Unio Retzius. (1788.) Simpson, 1900), p. 679 (restricted). Shell ovate, or more or less elongated, with straight longitudinal axis, not oblique, and beaks not very close to the anterior end. Beaks not very prominent, with shallow beak-cavities. Hinge-teeth well- developed. Outer surface without sculpture. Epidermis light or dark, with, or without, rays. Beak-sculpture distinctly of the double- looped type, or even zig-zag, with a distinct reéntering angle of the bars in front of the posterior angle. Often the sculpture is rudi- mentary, and consists of tubercles indicating the lower angles of the original loops. Soft parts much like those of Pleurobema, Elliptio, and Uniomerus. Mantle-connection between anal and supra-anal moderately long (generally almost as long as the anal). Inner lamina of inner gills free, except at anterior end. Marsupium formed by the outer gills, with the usual structure (see Plate XVIII, figs. 4, 5). Gravid females have not been seen by the writer, but the glochidia are described by European authors as being moderately large, subtriangular, with a hook on the ventral point of each valve. Type U. pictorum (Linneus). This genus chiefly differs from the foregoing genera in the shape of the glochidia and in the beak-sculpture. Although the marsupium is similar to the North American genera Pleurobema, Elliptio, and Uniomerus, | do not think that this indicates close relationship, but that it is due to parallelism of development. The genus Unio of the Old World has started from certain Unionine (with four gills serving as marsupium) in an independent line of descent. We do not yet know the forms which probably were ancestral to Unio. The shape of the glochidium indicates that somewhere near Unio was the starting point for the development of the subfamily Anodontine. bo ~I TS ANNALS OF THE CARNEGIE MUSEUM. Unio pictorum (Linnzus) 1758. See also Ortmann, I9QIICc, p. 21. A large number of specimens, both males and females, are at hand, from various parts of Germany and Hungary, received from W. Israél. Fic. 11. Unio pictorum (Linneus). Male, from Saale River, Rudolstadt, Germany. (Carn. Mus., No. 61, 4,934.) Branchial opening separated from the anal by a complete diaphragm formed only by the gills. Anal opening closed above by the union of the margins of the mantle, forming a supra-anal; this mantle-con- nection is rather long, slightly longer than the anal, and about as long as the supra-anal. Inner edge of bran- chial with distinct papilla, that of the anal almost smooth, or with very minute crenulations. In front of the branchial the inner edge of the mantle is practically smooth. Palpi subfalciform, their pos- terior, margins united for about one-third of their length, or slightly more. Gills (corresponding to the shape of the shell) rather long and narrow, the inner the wider, chiefly so anteriorly. Anterior end of the gills as usual, that of the inner widely distant from the palpi. Outer lamina of outer gill entirely connected mis ria Mere pillsiok aisreriic with the mantle; inner lamina of inner female, same locality. gill free from abdominal sac, with excep- tion of its anterior end. Both gills with well-developed water-tubes and continuous septa. The septa of the inner gill of the male (see Plate XVIII, fig. 3) and ORTMANN: FAMILIES AND GENERA OF NAJADES. 275 female (see Plate XVIII, fig. 4) are rather distant from each other; in the outer gill of the male, they are slightly more crowded. But in the female the septa of the outer gill are very close, forming narrow water-tubes (see Plate XVIII, fig. 4). The epithelial lining of the latter water-tubes is marsupial in character. Thus the marsupium is formed by the outer gills alone practically throughout their whole extent; for small sections at the anterior and posterior ends of the gill, which have wider water-tubes, pass gradually into the marsupial part, and may be disregarded. Gravid females are not at hand. According to previous observations of other authors (Harms, 1908, p. 696, fig. 1, and 1909, pp. 322 and 334; Haas, 1910a, p. 107), the glochidia are subtriangular, with hooks. Size 0.29 mm. Unio tumidus Retzius. The soft parts of four males and six females from Germany and Hungary are at hand, received from W. Israél. Structure essentially identical with that of U. pictorum. No gravid females have been observed. The glochidia have been figured by Schierholz (1889, pl. 4, fig. 63). Unio crassus Retzius.” Many specimens have been studied, received from W. Israél from the drainage of the river Elster in Thuringia, Germany. In this species also the soft parts are identical with those of U. pic- torum. The non-marsupial part at the anterior and posterior ends of the outer gills of the female is sometimes a little larger, but there are specimens: exactly like U. pictorum in this respect. Glochidia have not been observed, and have not been described so far as I know. Unio crassus musivus (Spengler).7! One male, and four females from Germany and Hungary have been sent to me by W. Israél. Absolutely identical in structure with U. crassus. A female from the river Begas, Hungary, had a few ovisacs near the middle of the 20 For nomenclature, see Thiele, 1909, p. 35. *1 This form is not batavus Maton and Rackett, 1907 (see Haas, 19104, p. 108, and I1910¢c, p. 167), but is surely batavus Lamarck, 1819, which name, consequently, cannot be used. As Haas (1910d, p. 62) has shown, the oldest name is musivus, Spengler, 1793. 276 ANNALS OF THE CARNEGIE MUSEUM. right outer gill filled with eggs. There were also eggs in the supra- branchial canal. No glochidia were seen. The water-tubes had no lateral water-tubes developed. Unio crassus consentaneus (Rossmessler). A male from the drainage of the Danube in Bavaria, and three males and four females from the Danube in Hungary, were sent to me by W. Israél. Agrees in all particulars with the foregoing forms. (A cross-section of the gills of the female is seen on Plate XVIII, fig. 5.) Genus PARREYSIA Conrad. (1853.)” Simpson, 1900), p. 840.—Ortmann, 1910), p. 139. Shell subovate or subquadrate, with rather high beaks, moderately deep beak-cavities, and well developed hinge-teeth. Epidermis bright, sometimes rayed. Beak-sculpture of the radial type: two sets of radial ridges run from in front and from behind the beaks in the direction of the lower margin. The two sets of radial ridges meet in the middle of the shell in an acute angle, and sometimes extend well upon the disk. Soft parts partly primitive, partly more advanced. Supra-anal separated from the anal by a well developed mantle-connection, which is rather long. Inner lamina of inner gills entirely connected with the abdominal sac. All four gills are marsupial in the female, with well developed septa and water-tubes, which latter are somewhat narrower in the outer gill thanin theinner. In the male, the septa are distinctly more distant than in the female. During pregnancy, the gills swell but little, and the edges remain sharp, and the ovisacs remain simple. Placentze subcylindrical, only slightly compressed, and not very solid. Glochidia not observed. This genus, in the structure of the soft parts, corresponds to Fus- conaja, Crenodonta, and Quadrula, to which it is apparently related, but represents another type of development of beak-sculpture, which may be derived from the simple Fusconaja-sculpture. Some minor features of the soft parts indicate that it has advanced a little along its * Determined only by the type-species, mullidentata Philippi =corrugatu Mueller (see Conrad, 1853, p. 267). The investigated species, wynegungaénsis Lea is closely allied to the type. ORTMANN: FAMILIES AND GENERA OF NAJADES. Arle own line, which is also indicated by the full development of the beak- sculpture. It may be possible, that species forming connecting links with Fusconaja still exist in eastern Asia. The investigation of ad- ditional, related types is much to be desired, and we should try especially to become acquainted with the glochidia. Parreysia wynegungaénsis (Lea). A number of specimens from Bombay, India, have been sent to me by L. S. Frierson. As to the description, I refer to my previous pub- lications (Ortmann, I910), p. 139, and 19IIa, p. 106, pl. 6, fig. 4, ply 7, hg: 3)- Genus LAMELLIDENS Simpson. (1900.) Simpson, 1900), p. 854.—Ortmann, I19IIa, p. 106. This genus bears about the same relation to Parreysia, as does Unio and Elliptio to Fusconaja and Quadrula. A complete diagnosis cannot be given at the present time, but the differences known to exist in the only species examined are the following: Outer gills alone marsupial, the shell more elongated, with the beak-sculpture rudi- mentary. As to the latter character, I may mention that I have seen, in a specimen of L. consobrinus, as well as in specimens of L. marginalis (Lamarck) in the Carnegie Museum, that the sculpture starts with a few (one or two) fine, concentric bars, and, following these, other bars are added, of which, however, only the lateral (anterior and posterior) parts are developed, which assume a direction radiating from the anterior and posterior side of the beak. These radiating ridges are very short, and I think they give us a clue as to the derivation of the radiating sculpture from the concentric. I think the Lamellidens- sculpture represents a phylogenetically older stage of beak-sculpture, while in other characters the form investigated is somewhat more advanced. Lamellidens consobrinus (Lea). One single sterile female from India has been investigated; I re- ceived it from L. S. Frierson. For the anatomy see my previous publication (Ortmann, I91Ia, p. 106, pl. 7, fig. 4). 278 ANNALS OF THE CARNEGIE MUSEUM. Subfamily A NODONTINA. To this subfamily belongs, first of all, the European genus Ano- donta, which is the typical genus, and which also occurs in North America, and probably likewise in Asia. In North America there are a number of additional genera, in some respects even more primitive than Anodonta, of which I have examined the following: Alasmidonta, Strophitus, Symphynota, Arcidens, Anodontoides, Lastena. They are all adopted from Simpson’s Synopsis, and I do not see any reason for changing these generic divisions. In the soft parts, they all very closely resemble each other. The fundamental idea, the physiological meaning of the anatomical pecu- liarities of this group, which governs its structure, is the following: these forms are bradytictic, and the breeding season becomes a long one, and the glochidia, after having fully developed, are not discharged, but kept in the marsupium over winter.** This makes necessary a special apparatus for supplying the glochidia with the necessary oxygen during this period. The problem is solved by the development of a special apparatus to secure the circulation of water within the gills, which, in the diagnosis (p. 224), has been called that of the “lateral water-tubes.”’ This apparatus exists only during the breeding season, but it has been found in all species the gravid females of which have been investigated. In sterile females traces of it are also generally discernible, since the lateral parts of the water-tubes often show indications of its presence in the conformation of their epithelium (see Plate XVIII, fig. 6). This is the most essential character of the subfamily. Other characters are furnished by the development of thickened tissue along the edge of the marsupium, which permits the distending of this gill during pregnancy, and this character is also generally easily seen in sterile females. Further, the mantle-connection sep- arating the anal and supra-anal is generally well, often very well, developed; the inner gill has the inner lamina free or connected with the abdominal sac. These latter two characters are of secondary value, but they help somewhat in the distinction of genera. The marsupium is always formed by the outer gills, the glochidia are rather large, subtriangular, and possess hooks. There are no generic distinc- tions observable in these characters, although the shape and size of 23 Very few Anodontine are known from countries without a winter, but such are present. It would be very interesting to study their behavior in this respect. ORTMANN: FAMILIES AND GENERA OF NAJADES. 279 the glochidia varies somewhat in the different species (see Plate XIX, figs. 2, 3, 4). In only one genus, Strophitus, the anodontine-structure of the marsupium has undergone a marked change, and has made a step in advance. Here each ovisac, which remains simple in other genera, is subdivided into a number of secondary compartments run- ning ina direction transverse to the gill. There are also further peculiarities in the marupium of this genus, which concern the mutual cohesion of the eggs and glochidia.** Strophitus is the only genus, which must be separated from the rest on solely anatomical grounds. In the case of all other genera the only criteria are the differences of the shells sometimes supplemented by minor characters of the soft parts. The shell, in this subfamily, is quite variable in shape, but in most forms it is not very thick, or it is even quite thin. The hinge is ex- tremely variable, showing all stages from a complete development to complete reduction. If present, the hinge-teeth are peculiar: the pseudocardinals are supplemented, in the lett valve by an additional (generally a third) tooth, which is formed by a projection of the interdentum, and extends below the corresponding part of the right valve. Sometimes this supplementary tooth is rudimentary, and it may be well separated from the posterior pseudocardinal, or may be fused with it. A very important feature of the shell seems to me the beak-sculpture, which indicates two, possibly three, lines of development within this sub-family. We would thus obtain the following scheme of classification: a1. Beak-sculpture double-looped, with a more or less sharp sinuation or re-entering angle. Mantle connection between anal and supra-anal moderate or very long. No tendency to unite the inner lamina of inner gills with abdominal sac. b}. Hinge-teeth more or less developed, at least pseudocardinals present. Shell not very thin. ci. Beak-sculpture not tubercular, and no sculpture upon the disk, but sometimes mpon’ the posterior slope..42-..-..0...--.4- Symphynota. c2. Beak-sculpture tubercular, and strong tubercles and other sculpture UWP OMUMeIGIS ewan te rieeeseys ors Siniens: Sen ORIN naoeeh voce wel sie cheraeler Arcidens. bo. Hinge-teeth completely absent. Shell smooth and thin......... Anodonta. ax. Beak-sculpture concentric, fine. Hinge-teeth practically absent. Mantle- 24 It is possible that the structural differences of Strophitus are connected with the fact recently discovered by Lefevre and Curtis (1911) that the glochidia of this genus do not pass through a parasitic stage on fishes. 280 ANNALS OF THE CARNEGIE MUSEUM. connection between anal and supra-anal moderately long. No tendency to unite the inner lamina of inner gills with abdominal sac.. . Anodontoides. a3. Beak-sculpture concentric, heavy. Mantle-connection between anal and supra- anal moderate. Certain forms with tendency to connect the inner lamina of inner gills with abdominal sac. bi. Hinge-teeth more or less developed, at least the pseudocardinals present. Marsupium with simple ovisacs. Shell subrhomboidal.. .Alasmidonta. bo. Hinge-teeth rudimentary. Marsupium of gravid female unknown. Shell Elongated. wii. Sucve oa ler che el eeomeupe ae Peabo bos ies alcay us naces eee eee Lastena. bs. Hinge-teeth rudimentary. Marsupium with ovisacs subdivided into trans- verse compartments. Shell subovate...................-. Strophitus. The most primitive types are undoubtedly Symphynota and Alas- midonta, and among them species with fully developed hinge-teeth are found. The most extreme modification of the soft parts is seen in Strophitus, while the most extreme specialization in the shell is represented by Anodonta (most successful adaptation to the life in quiet water with muddy bottom). Arcidens is peculiar in its shell sculpture. Anodontoides is a connecting form between a and 4s, but with the whole shell-structure more inclining toward aq. Lastena is yet rather doubtful in its position. It must be pointed out that comparatively few forms have been in- vestigated, and that further knowledge will possibly furnish the means for a better understanding ot the phylogeny of this subfamily. There surely should be Anodontine in Asia (aside from Anodonta proper), which possibly might be more primitive than any of those investigated hitherto. For the present, the most ancient types are known from North America, but I do not think that the subfamily originated in this continent. The shape of the glochidia indicates, that it started probably from a form near the European genus Unio, and this makes it more likely that the ancestral form lived in the Old World. Genus SYMPHYNOTA Lea. (1829.) (Simpson, 1900), p. 662.) Shell ovate or elliptic, compressed, with smooth disk, but sometimes with ribs upon the posterior slope. Beak-sculpture distinct, consisting of a few concentric bars, followed by others, which are distinctly sinuated,”®> or double-looped. Hinge with teeth, the pseudocardinals always present, the laterals present, imperfect, or absent. Soft parts of typical structure: outer gills alone marsupial, when 25 See description of beak sculpture of S. costata. ORTMANN: FAMILIES AND GENERA OF NAJADES. 281 charged their edges distending, lateral (secondary) water-tubes present, ovisacs not subdivided. Placentz very poorly developed, and only indicated when eggs are present. Inner lamina ot inner gills free from abdominal sac. Type S. compressa Lea. Very close to this stands S. viridis (Con- rad). These two species are normally hermaphrodites, while the other two (S. complanata (Barnes) and S. costata (Rafinesque)) are gono- chorists. This genus is the most primitive among the Anodontine with double- looped beak-sculpture. Simpson divides it into subgenera, which are well characterized, although there is not much need of a division of the genus on account of the small number of species. Symphynota compressa Lea. Numerous specimens from northwestern Pennsylvania and other parts have been investigated. This is a typical bradytictic form, and the breeding season is normal, beginning in August, and ending in May (and June in Lake Erie). The soft parts have been described by Lea (Obs., X, 1863, p. 423, as pressus), and Simpson (in Baker, 1898, p. 59). Anal and supra-anal separated by a well-developed mantle-connec- tion, which, however, is shorter than the anal. Inner edge of anal distinctly crenulated, that of the branchial with papilla; farther in front the edge is practically smooth, only in the beginning fine crenu- lations are seen. Palpi subfalciform, their posterior margins united for about one-half of their length. Gills broad, the inner the broader. Anterior end of inner gill about half way between the palpi and the anterior end of the outer gill. Gill-diaphragm normal. Inner lamina of inner gill free, except at anterior end. Gills with well-developed septa and water-tubes. This species being normally hermaphroditic (see Ortmann, 19110, p. 309), the gills have always (with extremely rare exceptions) the female structure, that is to say, in the inner gill the septa are rather distant, and the water- tubes are wide, and the outer gill is marsupial. When sterile, the septa are crowded, with marsupial epithelium, and the water-tubes are narrow. When gravid this gill swells considerably, and at the edge the tissue distends, so as to render the edge rounded off or trun- cated. Within this gill, each water-tube develops the characteristic 282 ANNALS OF THE CARNEGIE MUSEUM. lateral, or secondary, water-tubes, while the middle portion forms the ovisac, which is also closed at the base of the gill. The eggs fill the ovisacs in densely crowded masses, and in certain places a placenta- like cohesion may be observed. But when the glochidia are mature, they are perfectly free, and no indications of placentze are seen. Glochidia subtriangular, almost semicircular, longer than high, with hooks. Length 0.34; height 0.28 mm. (see: Lea, Obs., VI, 1858,° pl. 5, fig. 23; and Ortmann, 19110, pl. 89, fig. 10). Color of soft parts whitish, edge of mantle black, chiefly so poste- riorly. The foot is pale brownish yellow, the gills grayish. The ab- dominal sac is often pinkish. The charged marsupium varies greatly in color, this variation depending at least in part on the stage of de- velopment of the embryos. It may be white, cream-color, pinkish, pale orange, or various shades of brown. Symphynota viridis (Conrad). Numerous specimens have been investigated from the Potomac, Susquehanna, and Delaware drainages of eastern Pennsylvania. Breeding season from August to May. Soft parts described by Lea (Obs., XIII, 1874, p. 71). They are in every particular identical with those of S. compressa. This species also is hermaphroditic (Ortmann, 1911), p. 310), and specimens with the male structure of the gills have never been found. Glochidia (Lea, ibid., pl. 21, fig. 4) are about of the same shape as those of the fore- going species, but slightly larger. Length 0.36; height 0.30 mm. Color of soft parts as in compressa; marsupium cream-color, pale orange, or brown. Symphynota complanata (Barnes). Eight specimens from northwestern Pennsylvania, collected by my- self, have been investigated; in addition, three from the Kansas River, Lawrence, Douglas Co., Kansas (R. L. Moodie), and one from the Ohio at Portland, Meigs Co., Ohio (collected by myself). Among them were males, sterile and gravid females. Breeding season not completely known, but the dates at hand agree with those of other species. Eggs were found in the marsupium in September. Lea (Obs., X, 1863, p. 448) has described the soft parts; Simpson’s (in Baker, 1898, p. 61) description is partly incorrect (scalloped edge ~ ORTMANN: FAMILIES AND GENERA OF NAJADES. 283 of marsupium). A figure of the gravid female has been published by Lefevre and Curtis (1910, pl. 1, fig. 6), but the essential structure of the marsupium is not brought out. Mantle-connection between anal and supra-anal about as long as the anal, supra-anal slightly longer. In other respects, the soft parts are essentially identical with those of the other species of the genus. Glochidia of similar size, but shape more distinctly tri- angular, not so long in comparison with height. Length and height 0.34 mm. (see Lea, Obs., VI, 1858, pl. 5, fig. 29, and Lefevre and Curtis, l. c., p. 97, fig. A. The measurements given by Lefevre and Curtis, 0.29 X 0.30, are at variance with mine; see also Ortmann, I1911)b, pl dOp fig. 11): Color whitish when young, but foot and gills browner when old. Abdominal sac brown-orange, marsupium pale yellow to brown. Symphynota costata (Rafinesque). Many specimens from western Pennsylvania have been investigated, and two gravid females from Hurricane Creek, Gurley, Madison Co., Alabama (H. E. Wheeler, Sept. 13, 1910). Breeding season from August to May. Eggs were found only in August and September. Soft parts described by Lea (as Margaritana rugosa, Obs., X, 1863, p. 446) and Simpson (in Baker, 1898, p. 58). Soft parts like those of S. compressa. It is noteworthy that the anal is very large, and the mantle-connection between anal and supra- anal comparatively short, much shorter than the anal. Glochidia (see Lea, Obs., VI, 1858, pl. 5, fig. 26; Lefevre and Curtis, 1910, p. 97, fig. B, length 0.35; height 0.39), larger, more distinctly triangular, higher than long. Length 0.34; height 0.37 mm. This species is remarkable on account of its beak-sculpture, which does not conform to the double-looped type characteristic of this genus. In S. costata, the later bars (toward the disk) are rather heavy and straight, with hardly an indication of a sinuation. ‘The earliest bars are concentric as usual. But between the earliest and the latest, are some bars, which show an indication of sinuation more or less well-developed. This condition shows that we must not lay too much stress upon beak-sculpture as a general systematic character. S. costata is un- 284 ANNALS OF THE CARNEGIE MUSEUM. doubtedly a Symphynota, but the beak-sculpture is abnormally de- veloped. The sinuation of the bars, however, indicates that this species originally had double-looped sculpture. Color of soft parts rather remarkable. Orange tints are often found, similar to those seen in certain forms of Alasmidonta. The ground- color is yellowish brown; foot, margins of mantle, and adductors, often deep orange. The gills are brown, the edge of the mantle, as usual, blackish. The marsupium, when charged, varies from yellowish to brown. Genus ARCIDENS Simpson. (1900.) Simpson, 1900), p. 661. Shell subrhomboid, inflated, with full beaks. Disk sculptured. Beak-sculpture strong, distinctly double-looped, the loops tubercular, and the tubercles are continued in two radiating rows upon the disk. In addition, there are oblique folds upon the disk, and the posterior slope is also sculptured. Hinge with teeth, pseudocardinals present and well developed, laterals obliterated, but traces of them may be seen. Soft parts, as far as known, anodontine in structure, and similar to the genus Symphynota. Gravid females have not been observed. Type: A. confragosus (Say). The genus is incompletely known, but very probably it is to be placed near Symphynota. Arcidens confragosus (Say). I have one male and two females from Bayou Pierre, De Soto Parish, Louisiana, collected on Aug. 6, 1910, by L. S. Frierson, further the gills of an additional female from the same place, collected a little later, and one female from Pearl River, Jackson, Hinds Co., Mississippi, collected Nov. 5, 1910, by A. A. Hinkley. None of the females was gravid. The soft parts of the male have been described by Lea (Obs., X, 1863, p. 448). Supra-anal long, well separated from the anal by a mantle-connection, which is shorter than the anal. Inner edge of anal crenulated, inner edge of branchial with papillae. Diaphragm complete and normal. Inner lamina of inner gills free, except at ORTMANN: FAMILIES AND GENERA OF NAJADES. 285 anterior end. Posterior margins of palpi connected for not quite one-half of their length. Fic. 12. Arcidens confragosus (Say). Male from Bayou Pierre, De Soto Parish, Wa. (Carns Mus: No. 62, 4,701.) Gills anodontine in structure. Simpson (1900), p. 661) describes the marsupium in peculiar terms (‘‘of a peculiar, granular texture’’). Although I have not seen gravid females, the sterile females I possess offer nothing unusual or different from other Anodontine. Only the outer gills are marsu- pial, and their septa are much crowded, forming very narrow water-tubes, while in the inner gill the septa are much more distant. The septa of the outer gills are typically anodontine, and an indication of secondary water-tubes is present in the sterile female (see pla ec VITT, Fic. 12a. Left gills of a sterile fe- fig 6) Besides, at the edge of male, from Pearl River, Jackson, Hinds : ¢ : Co., Miss. Carn. Mus., No. 61, 4,928. the marsupial gill, there is a SING ue aa ante £928) thick mass of tissue, which indicates, that in the gravid female the edge is capable of distending. 286 ANNALS OF THE CARNEGIE MUSEUM. Genus ANoponTA Lamarck. (1799.) Simpson, 19008, p. 620. Shell elliptical, or elongated; thin; flat, or inflated, with smooth disk. Beak-sculpture distinct, but not very heavy, of the double- looped type, the loops separated by a sinuation, or a reéntering angle. Hinge-teeth completely absent. Only the outer gills are marsupial. When charged, the edge dis- tends, and secondary water-tubes are present. Ovisacs not sub- divided. No placentze are developed. Inner lamina of inner gills free from abdominal sac. Mantle-connection between anal and supra-anal generally very long, longer than either opening. Type A. cygnea (Linnezus). In the reduction of the hinge-teeth, in the long mantle-connection between anal and supra-anal, and the whole structure of the shell, this genus represents a very extreme specialization of the anodontine type. In the free inner lamina of the inner gill it is rather primi- tive. A large number of species are recognized by Simpson (19000), but in Europe the species-making in this group has gone beyond all the bounds of reason. A healthy reaction is, however, setting in, with regard to this genus as well as the European genus Unio (see Kobelt, 1908, p. 91; Thiele, 1909, p. 33; Israél, 1909, p. 26; Haas, 1910c). As will be shown below, the European genus Pseudanodonta, which has been split off, is also unsatisfactorily supported. It remains to be seen, whether the species from western North America and Asia have the same structure of the soft parts. Certain Chinese forms differ in the beak-sculpture. Anodonta cygnea (Linnzus). See also Ortmann, I9IIc, p. 22. A large number of specimens of both sexes, including gravid females, from various places in Germany and Hungary have been sent to me by W. Israél. I agree with Isra®l (1909) in regarding all Anodontas of central Europe (except complanata), as one species. The form cellensis is surely only the senile form of ponds, and anatina is the form of small creeks. The name of this species should be cygnea and not piscinalis. ORTMANN: FAMILIES AND GENERA OF NAJADES. 287 Supra-anal and anal openings widely separated, the one about as long as the other, but the united part of the margin of the mantle Fic. 13. Anodonta cygnea (Linneus). Male, from Obra South Canal, Sepno, Prov. Posen, Germany. (Carn. Mus., No. 61, 4,956.) between them longer than either, with slight variations in length. Anal with crenulations, branchial with papillae. Palpi with the pos- terior margins united for one- fourth or one-third of their length. Gills and diaphragm as usual, inner lamina of inner gills free, except at anterior end. In the male, the septa of both gills are rather distant; in the sterile female the septa of the outer gill ie are very crowded, forming very narrow water-tubes (see Plate XVIII, fig. 7). The whole outer gill is marsupial, and at its edge there is heavy tissue which permits the distending of the gill when charged. Within the marsupium Fic. 13a. Left gills of a sterile female, from Mogelnitza River, Prov. each water-tube is divided, in the z 3 Posen, Germany. (Carn. Mus., No. breeding season, into three tubes, Yen 61, 4,953-) two narrow lateral (secondary) water-tubes lying toward the faces of the gill, and a central larger 288 ANNALS OF THE CARNEGIE MUSEUM. tube; the latter forms the ovisac, containing the eggs and embryos, and this ovisac is also closed at the base of the marsupium by a fine membrane.”® The eggs and glochidia fill the ovisacs without forming placente, and the glochidia are discharged through the anal opening in rather irregular masses. Glochidia’? rather large, triangular, with hooks. They are about as long as high, 0.35 mm. (see Plate XIX, fig. 2). Having seen only alcoholic material I refrain from describing the colors of the soft parts. Anodonta complanata Rossmessler. See also Ortmann, I9IIc, p. 22. Six specimens from Dinkelsbueh], Bavaria, and ten specimens from Buda-Pest, Hungary, are at hand, received from W. Israél. Among them are gravid females. Fic. 14. Anodonta complanata Rossmessler. Male, from Weernitz River, Dinkelsbiihl, Bavaria. (Carn. Mus., No. 61, 4,958.) Inner edge of anal opening with very fine papilla. Gills (see Plate XVIII, fig. 9) essentially of the same structure as Anodonta cygnea, 26 That the lateral water-tubes are actually parts cut off from the original water- tube by folds is conclusively shown by fig. 8,-Plate XVIII. This is a slide made from a female, in which the eggs were just beginning to go into the marsupium. Attention should be called to the fact that in this species I never found the lateral water-tubes complete. This may be due to the fact that all my material was col- lected early in the breeding season. Nevertheless, some of the specimens had fully developed glochidia. 27 The glochidia have been figured by Fleming (1875, pl. 4, fig. 4) and Schierholz (1880, pl. 2. fig. 26), their measurements, 0.35 mm., have been given by Harms (1909, p. 332) and Haas (19104, p. 110). ORTMANN: FAMILIES AND GENERA OF NAJADES. 289 but septa more irregular in the non-marsupial gills. In the marsupial gill of the female, the septa are crowded, even more so than in A. cygnea. The tissue of the gills is generally more delicate in A. com- planata, but the various elements are similar to those of A. cygnea. The glochidia (see Plate XIX, fig. 3, also Schierholz, 1889, pl. 2, fig. 29; Fleming, 1875, pl. 3, fig. 11) are smaller, longer than high, with shorter hooks; thus the outline is less distinctly triangular, and less pointed. I find the length to be 0.34, the height 0.32; while Haas (1910a) gives 0.33 mm., but does not say in which dimension. For this species, Bourguignat (1880, pp. 11-13) has created the genus Pseudanodonta, founded originally upon the shape of the shell and differences in the hinge. The first character is quite pronounced, but cannot be regarded under any condition as a generic character; the second does not exist at all, which is best shown by the fact that it has been dropped entirely by subsequent authors (Haas). Other writers have added to the distinctive characters, which have been condensed by Haas (191t0a, p. 110; and 1910c, p. 170). According to Germain, the beak-sculpture is said to be different. Pseudanodonta is reported to have three to five tubercular ridges, which are absent in the true Anodonta, while in Anodonta, there ll) are flexuous ridges, but never tubercular ridges (“‘les Pseudanodontes ont . . . trois ll 4) mp | ) ) a cing rides tuberculeuses . . . qui man- quent chez les véritables Anodontes. Chez ——] oe ——) les Anodontes, les sommets sont parfois i —/ ornés de rides flexueuses, mais jamais de rides tuberculeuses’’). This statement isan intentional exaggeration of the actual con- wy : Fic. 14a. Left gills of a ditions, worded with the purpose to obscure sterile female from same lo- the similarities, and to emphasize the differ- cality. ences. The fact is that in both Anodonta and Pseudanodonta, the beak-sculpture is of the same type, and consists of a number of double-looped bars, of which, in complanata, the posterior loop is slightly more swollen, but not tubercular. Haas also described the beak-sculpture of Pseudanodonta as ‘‘ consisting of a few isolated, rather elevated tubercles,’’ which is positively wrong, as is shown by the specimens before me. 290 ANNALS OF THE CARNEGIE MUSEUM. According to Clessin, differences are said to be present in the gills. Clessin (1876, p. 446) asserts that the tissue of the gills in Pseudano- donta is more delicate, that the transverse striz (‘‘Querstreifen’’) are more deeply incised, and that the less conspicuous longitudinal striz (‘‘Leengsstreifen’’) are straighter and that the breeding compartments (‘‘Brutfecher’’) are more quadrate (‘‘bilden vollkommenere Quad- rate’’). The tissue of the gills is indeed more delicate, which is due chiefly to the slighter development of the interlaminar tissue. The gill- filaments are finer, and the interfilamentar grooves (probably the “‘Querstreifen”’ of Clessin) are deeper. What he calls ‘“‘ Lengsstreifen,’’ is probably produced by the longitudinal rows of water pores (ostia). Such a striation is indeed less distinct in a face view of the gills, in consequence of the stronger development of the filaments. They are not so distinctly visible as in A. cygnea, but nevertheless they are present. I cannot, however, under any conditions, see that they are straighter than in A. cygnea; on the contrary, they are slightly more irregular. What Clessin means by the statement that the ‘ Brut- feecher’’ are more quadrate, I cannot imagine. If he means the com- partments formed by the septa, I can only say that, in a face view, they are not quadrate at all, either in Anodonta or in Pseudanodonta; if he means in a cross-section, they are quadrate or approximately quadrate only in the outer gill of the sterile female, both in Pseudano- donta and Anodonta, while in all other gills, non-marsupial gills of the male and female, and marsupial gill of the gravid female, they are in both genera anything else but quadrate.*8 As has been said above, in Pseudanodonta the gill-structure is essentially the same as in the typical Anodonta, and all anatomical elements are present in both forms. The gills of A. complanata are indeed more delicate, chiefly in consequence of the slighter development of the interlaminar tissue; the gill filaments are finer, closer together, with deeper interfilamentary grooves, so that this layer on the outside of the gill is slightly thicker (compare Plate XVIII, figs. 7, 8, and 9). The consequence is that the rows of ostia, which are distinctly visible in A. cygnea in a face view, are rather indistinct in A. complanata. But when held up against the light, they also become distinct in the 28 Tt almost appears as if Clessin had compared the marsupial gill of a sterile female of Pseudanodonta, with a non-marsupial gill‘of Anodonta. But I am not sure of this, since his terms are rather vague. ORTMANN: FAMILIES AND GENERA OF NAJADES. 291 latter. Besides there is no difference whatever in the gills. The slight differences mentioned cannot be regarded, under any conditions, as of generic value, in face of the great similarity of structure. The presence of papillae on the anal opening (Haas, 1910a, p. 110) in Pseudanodonta is a character, which at most has merely specific value. There remains only the glochidium (see Plate XIX, fig. 3) to be considered. There are, indeed, certain differences between the glochidia of A. complanata and cygnea, but as far as they are known in other members of the genus and subfamily, such differences are encountered elsewhere, without being considered as of generic value. The glochidia are undoubtedly built upon the same plan in both species. A. complanata not only is a true Anodonta, but judging by the shape of the beak and beak-sculpture belongs to that group in the genus, of which A. cygnea is the type. We cannot separate it generically from the latter, without disregarding natural affinities. Anodonta imbecillis Say. I have investigated twenty-three specimens from northwestern Pennsylvania (Allegheny and Lake Erie drainages), and two from Lawrence, Douglas Co., Kansas (R. L. Moodie). All were females in structure, and many were gravid. This species is hermaphroditic. Typically bradytictic, and gravid from September to May. In Lake Erie the time of discharging the glochidia is postponed even further, this act having been observed as late as July 12. The anatomy has been described by Lea (Obs., X, 1863, p. 449). Of all American forms this species resembles most closely the European Anodontas in the shape of the shell. It differs, however, in being hermaphroditic. The soft parts present no special features, and they agree both with the European A. cygnea and the: North American A. grandis. The anal opening is almost smooth. Glochidia smaller than those of cygnea, slightly higher than long (length 0.30; height 0.31 mm.), of the usual shape, with hooks (see ea Obs, Viv 1858; pl. 5, fig. 360; Ortmann, 1O11d, pl: 890,. fig. 13). Color of soft parts whitish, foot yellowish brown; gills pale brown; edge of mantle blackish. Charged marsupium brown. Anodonta henryana Lea. Seven specimens from a branch of the Rio Grande, Mercedes, Hidalgo Co., Texas, collected May 15, 1907, by Dr. D. A. Atkinson. 292 ANNALS OF THE CARNEGIE MUSEUM. Soft parts and glochidia described by Lea (Obs., VIII, 1860, p. 373). I think my specimens belong to this species, although they are all remarkably shortened and truncated posteriorly. They are all gravid, with glochidia. Possibly this species also is hermaphroditic. Soft parts absolutely identical with those of A. imbecillis, and the glochidia also are practically identical. Length 0.29; height 0.30 mm. (The difference from the measurements given for A. imbecillis may be due to the personal equation of the observer.) The shell of this species reveals that it undoubtedly belongs to the same group as A. imbecillis. Anodonta grandis Say. Numerous specimens of the typical form as well as of several vari- eties have been investigated. They come from western Pennsylvania, . Kansas, northern Alabama, and Louisiana. The breeding season begins in August and September, and ends very early in spring, for in April the majority of the females have already discharged their glochidia. But occasionally gravid females are found as late as the end of May (latest date, May 22). The soft parts have been described (as of A. plana, decora, ovata) by Lea (Obs., X, 1863, p. 452) and by Simpson (in Baker, 1898, p. 52, chiefly as to color). Mantle-connection between anal and supra-anal very long, anal small, its inner edge crenulated. Branchial opening with papille, inner edge in front of branchial smooth. Posterior margins of palpi con- nected for one-third to one-half of their length. Anterior end of inner gills midway between that of the outer gills and the palpi, so that it is widely remote from the palpi. Diaphragm normal. Inner lamina of inner gills free except at anterior end. Septa of the gills distant in the male and the non-marsupial gills of the female. Marsupium formed by the outer gills, which swell greatly when charged, distending at the edges so as to appear truncated, and developing lateral water- tubes, while the ova are contained in the central ovisacs; the latter are not subdivided. No placente are formed, and the glochidia are discharged loose, in irregular masses. Glochidia very large, the largest known to me, even larger than those of A. cygnea, subtriangular, slightly higher than long (length 0.36; height 0.37 mm.) with hooks. The glochidia figured by Lea (Obs., VI, 1858, pl. 5, figs. 32-34) as of A. lewisi, ovata, decora, differ some- what from each other, while they actually should be all alike. ORTMANN: FAMILIES AND GENERA OF NAJADES. 293 Color whitish. Foot orange-yellow, palpi and gills brown. Edge of mantle brown, black behind. Charged marsupium yellowish white (with eggs) to liver-brown (with glochidia).22 The orange tint of the foot is variable, lighter or darker. Anodonta cataracta Say. About twenty-five specimens from various places in the Atlantic drainage of eastern Pennsylvania have been investigated. The breeding season begins in August, in which month I repeatedly found gravid females with eggs. I have no other observations of my own, but Conner (1907, p. 88) gives October to May as the breeding season. Lea (Obs., II, 1838, pl. 15, fig. 46) has figured the animal, but his figure is practically useless. The soft parts resemble in every respect those of A. grandis. I have seen the glochidia of specimens from the Delaware River, collected by C. H. Conner on March 19, 1911, and they agree with the figure published by Lefevre and Curtis (1910, p. 97, fig. C); the dimensions are: length 0.36; height 0.37 (identical with those of the glochidia of A. grandis). The colors of the soft parts are also the same. I have my doubts as to the specific distinctness of this form from A. grandis. At any rate, it is merely the eastern representative of the western grandis, and does not have any close affinity to the European species and the Anodontas of the Pacific slope of America, as Walker (1910a, p. 135) believes, and there is no reason to think that ‘it was a co-immigrant with Margaritana margaritifera to the east coast of North America” (from Europe). A. cataracta is, if anything, an east- ern offshoot of the A. grandis-stock of the central basin. Genus ANODONTOIDES Simpson. (1898.) (Simpson, 19008, p. 658.) Shell subelliptical, thin, inflated, with smooth disk. Beak-sculpture distinct, but not very heavy, consisting of concentric ridges curved up behind, not double-looped. Hinge-teeth absent, or represented by the merest rudiments. Soft parts much like those of Anodonta. Outer gills alone marsupial, when charged distending at the edges, and secondary water-tubes present, ovisacs not subdivided. No placente developed. Inner 22 Sometimes the light color is preserved in the glochidial-stage. 294 ANNALS OF THE CARNEGIE MUSEUM. lamina of inner gills free from abdominal sac. The mantle-connection separating anal and supra-anal shorter than in Anodonta. Type A. ferussacianus (Lea). This is practically an Anodonta with concentric beak-sculpture. The general make-up of the shell is much like Anodonta, while the beak-sculpture is suggestive of the Alasmidonta-group, without being so heavy. Anodontoides ferussacianus (Lea). Numerous specimens of the typical form, as well as of the var. subcylindraceus (Lea) have been investigated, the former from the Ohio drainage in western Pennsylvania, and the Cumberland River in Kentucky, the latter from Lake Erie. Bradytictic. The breeding season begins in August and ends in May; discharging specimens were found on May 14. Soft parts described by Lea (Obs., X, 1863, pp..449 and 451), and Simpson (in Baker, 1898, p. 73), but the latter is mistaken with reference to the marsupium of ferussacianus, while the description of that of subcylindraceus (p. 74) is correct. Anatomy essentially that of Anodonta, but it should be mentioned that the mantle-connection between anal and supra-anal is only about as long as the anal as well as the supra-anal. The inner edge of the anal is finely, but distinctly papillose. The posterior margins of the palpi are only connected for a short distance, and the anterior end of the inner gill is about half-way between that of the outer gills and the palpi. The marsupium has the same structure as in Anodonta, and the glochidia (Ortmann, 19110, pl. 89, fig. 12) are rather small for the subfamily, subtriangular, and about as long as high (0.32 mm.). They have hooks, although Lea (Obs., VI, 1858, pl. 5, fig. 35) figures and describes them as without hooks. I find that by a singular oversight I failed to make any field-notes on the color of the soft parts of this species but from alcoholic material and according to my recollection it is grayish white with the foot and the gills inclining to brownish. The marsupium containing glochidia is brown. Genus ALASMIDONTA Say. (1818.) (Simpson, 1900), p. 666.) Shell elliptical, or generally rhomboidal, inflated, with a well- developed posterior ridge. Disk generally smooth, but sometimes ORTMANN: FAMILIES AND GENERA OF NAJADES. 295 with a faint sculpture upon the posterior slope. Beak-sculpture heavy and coarse, the later bars are often very thick and swollen, concentric, often angled behind, not double-looped. Hinge with teeth, pseudocardinals always present, laterals present or absent; in the former case sometimes abnormally developed. Only the outer gills are marsupial, when charged, distending at edges, secondary water-tubes present, and ovisacs not subdivided. No placentz developed. Inner lamina of inner gills free from ab- dominal sac, or more or less connected with it. Mantle-connection between anal and supra-anal not very long. Type A. undulata (Say). This genus is rather primitive, especially in the character of the shell, and stands on about the same level as Symphynota, representing an- other parallel branch, characterized by the heavy, concentric beak- sculpture. The tendency to a union of the inner lamina of the inner gills with the abdominal sac indicates a slight advance in structure. Alasmidonta heterodon (Lea). About thirty specimens collected in April near Philadelphia have been investigated. Among them were many gravid females. Conner (1909, p. 112) found this species gravid in February. An imperfect description of the marsupium and the glochidia has been given by Lea (Obs., X, 1863, p. 442). The soft parts do not offer anything remarkable. The mantle- connection between anal and supra-anal is shorter than the anal. Anal with crenulated inner edge. Inner lamina of inner gills free. Posterior margins of palpi connected for about one-half of their length. The anterior end of the inner gill is separated from the palpi, but nearer to them than to the anterior end of the outer gill, Marsupium typically anodontine in structure. Glochidia (Ortmann, tort), pl. 89, fig. 8) the smallest known to me in this subfamily. They are subtriangular, much longer than high, with strong hooks. Length 0.30; height 0.25 mm. Color of soft parts whitish; charged marsupium brown. This is in every respect the most primitive type known to me in this subfamily. Alasmidonta minor (Lea). One male, and one gravid female (with glochidia), from Cumberland River, Pineville, Bell Co., Kentucky, have been communicated to me by B. Walker. 296 ANNALS OF THE CARNEGIE MUSEUM. Of this form I have only the soft parts, and their structure is like that of other species of this genus. The mantle-connection between anal and supra-anal is almost as long as the anal, and the supra-anal is only slightly longer than the mantle-connection. The anal is finely crenulated, and the branchial has papilla. Posterior margins of palpi connected for a short distance. Inner lamina of inner gills free. Anterior end of inner gill about half-way between the palpi and the anterior end of the outer gill. Marsupium as usual, with distended edges, and secondary water- tubes. Glochidia as small as in the foregoing species, and of the same shape. Length 0.30; height 0.25 mm. (see Plate XIX, fig. 4). Color whitish, edge of mantle spotted with black and white in the region of the branchial, anal, and supra-anal. Marsupium brown. I have had no opportunity to examine the shell of this species, but it seems that it is related to A. heterodon. Alasmidonta undulata (Say). Numerous specimens from the Atlantic drainage in eastern Penn- sylvania have been in my hands. The breeding season begins in the middle of July, and lasts till the middle of June, so that the end of one season, and the beginning of the next are not very far apart. Of specimens found in July, 18th and 22d, all had only eggs; while those found on June 14 (only two) had fully developed glochidia. Soft parts typical. Mantle-connection between anal and supra-anal rather long, slightly longer than the anal and than the supra-anal. Branchial with papilla, anal crenulated. Posterior margins of palpi connected for one-half or slightly less than one-half of their length. Diaphragm normal. Inner lamina of inner gills entirely connected with the abdominal sac, and only in a few cases was a small hole ob- served at the posterior end of the foot. Anterior end of inner gill about half-way between the outer gill and the palpi. Marsupium normal; when charged, having distended edges, secon- dary water-tubes, and undivided ovisacs. Glochidia (Ortmann, 1911d, pl. 89, fig. 9) moderately large, higher than long, with strong hooks. Length 0.34; height 0.36 mm. Abdominal sac whitish; foot paler or darker orange-brown; palpi whitish to orange-brown. Gills grayish brown, shading into orange. Mantle transparent gray, shading into brownish or brownish orange ORTMANN: FAMILIES AND GENERA OF NAJADES. 297 on the margin anteriorly, into white posteriorly. Edge of mantle brown, posteriorly orange, mottled with black spots. Adductors grayish to orange. The orange tints are often rather pale, inclining toward grayish yellow. Marsupium charged with eggs pale yellow; with glochidia brownish. Alasmidonta marginata (Say). Of this species, and of its eastern variety varicosa (Lamarck) a large number of specimens have been investigated, both from the Ohio and the Atlantic drainages in Pennsylvania. Bradytictic, breeding season beginning in August, and lasting until May. Discharging specimens have been found on May 3. The anatomy has been discussed by Lea (Obs., X, 1863, p. 446) and Simpson (in Baker, 1898, p. 63). Soft parts essentially like those of A. undulata, to which species it is indeed closely allied. The mantle-connection between anal and supra- anal is much shorter. I have found the inner lamina of the inner gills always connected with the abdominal sac, although Lea (/. c.) says that it is sometimes more or less free at the posterior end. Posterior margins of palpi connected at base only. Glochidia (see Lea, Obs., VI, 1858, pl. 5, fig. 27) rather large, higher than long, with hooks. Length 0.33; height 0.36 mm. Color entirely like that of A. undulata, with a strong tendency toward orange tints; posterior margin of the mantle spotted with black and orange. Marsupium, according to contents, yellowish white to brown. Genus LASTENA Rafinesque. (1820.) (Simpson, 1900), p. 654.) Shell elongated; not inflated; without distinct posterior ridge. Disk smooth. Beak-sculpture concentric, bars irregular, coarse, middle part nearly straight. Hinge with rudimentary teeth, pseudocardinals only vestigial, laterals absent. Soft parts only of male and sterile female known, but as far as can be seen anodontine in structure. Inner lamina of inner gills free. Mantle-connection between anal and supra-anal rather short. Type L. lata (Rafinesque). According to the characters of the shell, this genus stands between 298 ANNALS OF THE CARNEGIE MUSEUM. Alasmidonta and Strophitus. It has the beak-sculpture® of the genera of the Alasmidonta-group, and approaches Strophitus in the hinge. It is very much to be regretted that gravid females are not at hand, and that the structure of the charged marsupium and the glochidia remain unknown. The sterile females, which I have seen, make it clear that only the outer gills are used as marsupia, and that they havea structure like that found in sterile females of the Anodontine in general. Lastena lata (Rafinesque). I have received, from B. Walker, the soft parts of two males and two sterile females from the Cumberland River in Pulaski and Cumber- land Cos., Kentucky. Anal and supra-anal separated by a rather short connection of the mantle. Anal finely crenulated, branchial with papillae. Posterior margins of palpi connected for a short distance. Gills and gill-diaphragm normal, and not as described by Simpson,* Gills long and rather narrow, the inner one decidedly wider in front, its anterior end distinctly in front and below the anterior end of the outer gill, but separated from the palpi by a short, but distinct interval (it is connected with the descending part of the mantle attachment line for about three-fourths of its length, while one-fourth is occupied by the interval). Posteriorly, the gills do not project freely, but are entirely of the usual shape. Inner lamina of inner gills free from the abdominal sac with exception of the anterior end. Septa and water-tubes normally developed. Marsupium formed by the outer gills, with the water-tubes narrow, and the septa close together, thick, and with strongly wrinkled epithelium. In the females at hand, no indications of secondary water-tubes could be seen. The thickened tissue at the edge of the gill was also not well developed. Both specimens are small (under medium size, about 40 and 45 mm. long), and seem never to have been gravid. Color (of alcoholic material) whitish, edge of mantle brown, black behind. 30 This sculpture is somewhat variable. It begins with simple concentric bars upon which a few heavier bars follow, slightly sinuated in the middle and angled behind, and then follow a few more upon the disk, quite heavy, but indistinct. The latter are sometimes absent, and sometimes even the sinuated bars are very rudimentary. The sculpture resembles somewhat that of Strophitus. 31 Simpson (1900), p. 654) says: “inner and outer (gill) about alike in size and form, projecting free slightly behind.” ORTMANN: FAMILIES AND GENERA OF NAJADES. 299 Genus STROPHITUS Rafinesque. (1820.) (Simpson, 1900), p. 616.) Shell subelliptical, subovate, or subrhomboidal, inflated, with indis- tinct posterior ridge. Disk smooth. Beak-sculpture concentric, con- sisting of a few rather heavy bars curving sharply up behind, forming an angle there. Hinge-teeth quite rudimentary, only mere vestiges of pseudocardinals present, which sometimes disappear altogether. Inner lamina of inner gills free, or more or less connected. Mantle- connection between anal and supra-anal rather short. Marsupium formed by outer gills, when charged having the edge distended and secondary water-tubes. But the ovisacs do not remain simple, and are subdivided into a number of compartments running cross-wise in the gill from face to face; each compartment containing the ova and glochidia is well defined; placentule solid, persistent until they are discharged. Type S. undulatus (Say). This genus offers in the marsupial structure the highest specialization known among the Anodontine. In the hinge and the inner lamina of the inner gill we also see indications of a high stage of development. The beak-sculpture and other characters of the shell assign it a place in the Alasmidonta-series. Strophitus edentulus (Say). I have examined a great many specimens from all over Pennsylvania, from the Ohio, as well as the Lake Erie, Delaware, Susquehanna, and Potomac drainages. I have also seen specimens from the Erie drainage in Huron Co., Ohio (O. E. Jennings coll.), from the Potomac drainage in Maryland (collected by myself), from Lawrence, Douglas Co., Kansas (R. L. Moodie), and of the form called shafferiana Lea, from the Cumberland River in Kentucky (B. Walker). Bradytictic. The breeding season begins in July (earliest date July 11), and ends in April and May. Discharging specimens have 82 It is not clear what the original Anodonta undulata of Say is. Most authors (including Simpson) have taken the common Strophitus of the Atlantic drainage for it; but this is not different from the western edentulus Say. Conner thinks he has re-discovered the real undulatus in the tide waters of the Delaware river, but I can only see a local form of edentulus in it (I have not seen its soft parts). The form I have investigated is surely the edentulus of Say. 3800 ANNALS OF THE CARNEGIE MuSEUM been found on April 22 and 24, and May 6 and 11. The latest date for gravid females in spring is May 22.8 Lea (Obs., I], 1838, pl. 15, fig. 47) has published a very poor figure of the soft parts, and descriptions of the latter (Obs., X, 1863, pp. 450 and 453). Mantle-connection rather short, shorter than the anal. Inner edge of anal crenulated, that of branchial papillose. Posterior margins of palpi connected at base only. Diaphragm normal. Inner lamina of inner gills very variable. In most cases it is connected anteriorly for about one-half of the length of the abdominal sac,”in other cases the connection is much longer, and not unfrequently the inner lamina is entirely connected. Anterior end of inner gill about halfway between that of the outer gill and the palpi. Septa of the gills distant from each other in the male and the non-marsupial gills of the female. The outer gill alone is marsupial, with much more crowded septa. When gravid, the gills swell greatly, distend at the edges, and lateral water-canals are formed. In addition, the ovisacs are divided into compartments as described above. Pla- centule containing two to ten (or more) ova or glochidia. The placentule and glochidia have been figured by Lea (VI, 1858, pl. 5, figs. 37 and 38). The latter are not very large, subtriangular, longer than high, and have hooks. Length 0.36; height 0.30 mm. Color of soft parts quite variable, but with the same tendency as Alasmidonta undulata to have certain parts (foot, edge of mantle, and adductors) orange. Some specimens (chiefly young ones) are more or less uniformly grayish or yellowish white, while others exhibit all shades from yellowish through brownish orange to bright orange. Gills mostly paler or darker brown, shading to orange. Mantle edge blackish, with the same orange and black spots as Alasmidonta un- dulata. Marsupium, according to contents, pale yellow, or creamy, to brown. Subfamily LAM PSILINA. A large number of forms belong here, which, as far as we know, are all found in North America, extending southward into Central America. Indeed, Simpson associates with these a number of Asiatic and African 33 According to Lefevre and Curtis (1911) the “‘interim”’ is in July in Wisconsin, probably corresponding to the higher latitude. In 1911 I found gravid and dis- charging females in West Virginia on May 11, 12, and 13, but on May 23, 24, and 25 no gravid females were any more seen. ORTMANN: FAMILIES AND GENERA OF NAJADES. 301 genera (Pseudospatha, Hyriopsis, Chamberlainia, Cristaria, Lepido- desma, Pilsbryoconcha), but of all these the soft parts are unknown, and we may entertain strong doubts as to their belonging here. In order to understand the structure of the Lampsiline, and their further differentiation, we must recall their essential characteristics and their purpose. We have seen that the soft parts are accommo- dated to two functions: (1) owing to the extended breeding season (in bradytictic forms) that of securing the proper aération of the gravid marsupium, (2) the discharge of the glochidia through the edge of the marsupium. The latter physiological character is unique, and is found only in this subfamily. The former occurs in the Anodontine, but we have seen that it is there brought about in another way. Very likely the adaptations to these two peculiar functions are con- nected in a degree. We see that it is the general tendency among the Lampsiling to move the marsupium toward the edge of the gill, and even beyond the latter. This has the effect that it is removed, more or less, from the natural outlets, and comes in close contact with the outer water flowing over the gills. Under these conditions it is easily understood that the habit was acquired to discharge the glochidia not by the long way (the suprabranchial canals), but by the shortest, by making them go through holes in the edge of the marsupium (see Plate XVIII, fig. 10). Thus we may say that the lampsiline mar- supium serves two purposes, and is built according to a type which meets first the necessity of aérating the marsupium, and which in con- sequence of the structure so assumed, made another peculiar way of discharge desirable. To supply breathing water for the glochidia, however, is of chief importance, and thus the further differentiation within this subfamily is easily understood, when keeping this point in view. Allowance should be made for certain expressions used in the following statement, and they should be excused by my desire to make the facts as clear as possible. Among the Lampsiline, there are at least four types of marsupial structure, which represent as many different attempts to find a way of supplying breathing water to the marsupium. They all agree in having as a common feature the extrusion of the marsupium beyond the edge of the gill, and its investiture by only a very thin membrane, so that osmotic processes are greatly facilitated. In addition, in three of these types, there is also developed the tendency to move the 302 ANNALS OF THE CARNEGIE MUSEUM. marsupium toward the posterior part of the gills and the shell, in order to have it as close as possible to the branchial opening and the inflowing water. Of the four types of marsupium, three are found only in compar- atively a few forms, while the fourth is more widely distributed, and gives origin to a new line of development. The first three may be called rather indifferent attempts on the part of the forms concerned, to solve the problem. The problem has been solved by them, indeed, but the way in which they did it did not contain any further pos- sibilities. In the fourth case, the attempt was more successful, and opened the way for a series of additional improvements. 1. In one case (Ptychobranchus), the marsupium remains in a primitive stage in this respect, that it is pushed only slightly beyond the edge of the gill, and is not moved backward, but occupies the whole gill, But here in order to insure proper aération by increasing the surface of the marsupium, while the latter remains rather thin, the whole marsupium is thrown into a number of folds which permit the water to easily reach the ovisacs, which are subcylindrical and not much swollen. 2. In the second type (Obliquaria and Cyprogenia), the task has been accomplished by reducing the number of ovisacs. This would have had the result of restricting the number of ova that could be accom- modated in the marsupium, but this disadvantage is counterbalanced by a tendency to greatly elongate the ovisacs, in the direction beyond the edge of the gill. This feature is only slightly developed in Obli- quaria, while it reaches its greatest perfection in Cyprogenia, and here there is not room enough within the shell for the extremely elongated ovisacs and thus they have to coil up in a spiral. 3. In the third type (Dromus), the marsupium originally is rather simple, the ovisacs remaining subcylindrical or being only slightly compressed. Here a better aération is accomplished by a peculiar arrangement of the glochidia within each ovisac. They are not dis- tributed through the mass of the placenta, but are situated along the edge of the slightly compressed placenta, thus facing the outer walls of the marsupium, where they are nearer the breathing water. In addition Dromus has developed a peculiar warping and folding of the marsupium, which also apparently has the object of increasing the surface offered to the water. But this latter feature is exhibited only in old specimens. it reminds somewhat of the structure of Ptycho- ORTMANN: FAMILIES AND GENERA OF NAJADES. 303 branchus, but I think it has been independently acquired, and does not indicate close relationship. 4. In the fourth type (all other genera), an entirely different ar- rangement to provide aération for the marsupium has been effected. While in the first three types structural modifications of or within the marsupium are introduced to bring the glochidia close to the breathing water, here the marsupium itself remains rather simple, and it is the water supply which is increased and intensified. The marsupium forms in this case a rather swollen, generally kidney- shaped mass, in which the ovisacs are transversely dilated, so as to give them a leaf-like shape. The tendency to locate the marsupium in the posterior part of the gill and to cause it thus to approach the posterior end of the shell, close to the branchial opening and close to the incoming water, is common to all these genera. In other respects, there is no further differentiation of the marsupium. Buta new device begins to develop, having for its aim the increase of the flow of the water over the marsupium, and this is accomplished by special struc- tures on the edge of the mantle, just in front of the branchial opening. In the simplest cases (Obovaria, Nephronajas, Amygdalonajas, Pla- giola, Paraptera, Proptera), the inner edge of the mantle is only slightly dilated, forming a fine lamella, but the presence of a (muscular) thickening, and often of pigment, indicates, that the edge has here a peculiar function. In other genera (Medionidus, Eurynia, Lamp- silis), this part of the inner edge of the mantle is greatly developed, and carries special appendages in the shape of strong papille or flaps, which have the function of producing by their contractions, a lively current of water over the surface of the marsupium, which lies immedi- ately inside of them. Finally, in the genus Truncilla, the inner edge of the mantle, which also has papille, is removed from the outer edge, thus enclosing a separate compartment, which may possibly be regarded as a kind of reservoir. Thus it becomes evident that of these four arrangements acquired by the Lampsiline to provide breathing water for the glochidia, the last was the most advantageous, because it included the possibility of further development and improvements such as we find realized in the various genera just mentioned. 5. There remains yet a fifth type of structure, that found in Frier- sonia. Here the marsupium is truly lampsiline, resembling somewhat the Obovaria-type, but it is not so swollen, and instead of being blunt, 304 ANNALS OF THE CARNEGIE MUSEUM. the edge is here sharp. The water-tubes (ovisacs) have a peculiar curve backward toward a point near the posterior end of the mar- supium. For the present, I cannot correlate this structure with any special function, and must leave it for future study to decide what the meaning of this feature is. We may arrange the genera of this subfamily as follows: ai. Marsupium thin, not kidney-shaped, ovisacs subcylindrical or very slightly compressed. Placentz generally rather solid. Inner edge of mantle in front of branchial opening not distinctly differentiated. Shell rounded, ovate, or subelliptical, sometimes with outer sculpture. Male and female shells practically alike. hi. Marsupium occupying the whole of the edge of the outer gill, folded. Pla- cente subcylindrical, club-shaped, short, Shell subelliptical. Ptychobranchus. b2. Marsupium occupying only a part of the outer gill. Placenta moderately elongated or very long, subcylindrical, or very slightly compressed. Shell more or less rounded, generally with tubercles. a. Marsupium consisting of comparatively few, elongated ovisacs, ex- tending from near the base of the gill beyond its edge. Placente subsolid, subcylindrical. di. Placente moderately long, slightly curved. Marsupium just behind theymuiddlesonthe (gillian re ee Obliquaria. dz. Placente very long, spirally coiled up. Marsupium in, or slightly inmront of they middlerorf theres ere eee Cyprogenia. ¢. Marsupium consisting of a large number of ovisacs, occupying the larger posterior section of the outer gill. Placentz slightly com- pressed near base, tapering, and becoming subcylindrical toward the margin. d;. Ovisacs straight. Placente almost entirely beyond the edge of the gill, subsolid. _Glochidia much longer than high, arranged on the outer faces of the marsupium. Edge of marsupium blunt, in old specimens warped and folded. Shell rounded, disk nodular. Dromus. dz. Ovisacs strongly curved back in the distal part. Placente for a great part enclosed between the original gill lamina, not solid. Glochidia not longer than high, distributed uniformly through the placental mass. Edge of marsupium sharp, pointed behind, not folded. Shell subelliptical, disk smooth............. Friersonia. a2, Marsupium thick, more or less kidney-shaped, ovisacs dilated and compressed. Placentze not very solid. Inner edge of mantle in front of branchial opening, more or less differentiated. Shell rounded, elliptical, or elon- gated, swollen or compressed, generally without any outer sculpture. Male and female shells more or less different in shape. by. Inner edge of mantle slightly lamellate and crenulated, but without dis- tinct papilla or flaps. Male and female shell differing only slightly in shape, sometimes hardly at all. ORTMANN: FAMILIES AND GENERA OF NAJADES. 305 ci. Shell rounded, ovate, or subelliptical, without or with indistinct posterior ridge. Glochidia of normal size and shape, subovate. d,. Shell rounded or ovate, swollen. Epidermis brownish, rarely greenish, Valen, siaahGunics IghebcooeecapalaoouacecHoOnecCoaaUlé Obovaria. ds. Shell subovate or subelliptical, compressed, or only slightly swollen. Epidermis greenish or yellowish, with more or less distinct rays. Nephronajas. c.. Shell ovate, triangular, swollen, or subelliptical and compressed. Glo- chidia of abnormal size or shape. dy. Shell subovate or subtriangular, with a strong posterior ridge. e1. Glochidia of normal shape, but abnormally small size. Shell SuboOVvateronelongates. a-racitetas cles er cke cries Amygdalonajas. ex. Glochidia spatulate, with gaping margins, large. Shell sub- triam ouilary peer iacereta crerecclauere ero ahd wlio meus eh dorian cba Srelovs\eite ey Plagiola. d2. Shell subovate or subelliptical, more or less compressed, often winged, without distinct posterior ridge. e1. Glochidia of normal shape, but abnormally small. Shell rather (ELON ays aehete te sh ee dere he eta tee ld he aaa Cmte eA ae Paraptera. ex. Glochidia celt-shaped, with two spines on each valve. Shell thin OL rath erathi Ghent spacysty don sporte las erect orale stelefalorsc Proptera. bs. Inner edge of mantle with papillz or flaps. Male and female shell distinctly, and often greatly different in shape. ci. Inner edge of mantle parallel with and close to the outer edge. Shell ovate, elliptical, or elongated. Glochidia subovate. di. Inner edge of mantle with papille. e1. Shell with nodulous plications upon the posterior slope. Medionidus. exmsnelliwithoute Sculpture sarierciem cisetcisiotie: oii eter ete Eurynia. d». Inner edge of mantle forming a ribbon-like flap......... Lampsilis. co. Inner edge of mantle in front of the branchial opening more or less remote from outer edge in the female. Shell variable, not ovate or elliptical. Glochidiaalmostsemicirculatsaccrcciee cece iia cers oe Truncilla. In all these genera, we have a beak-sculpture, which is rather rudi- mentary, and, when developed, either of the concentric or the double- looped type. Beak-sculpture in this subfamily is apparently a char- acter becoming more or less obliterated, and thus cannot be used for general systematic purposes, although it is available as a subsidiary character in a few cases. Genus PTyCHOBRANCHUS Simpson. (1900.) (Simpson, 19000, p. 612.) Shell subelliptical, somewhat elongated. Disk smooth, sometimes with ridges on the posterior slope. Beak-sculpture indistinct, con- sisting of a few ridges, the first concentric, the others slightly double- 306 ANNALS OF THE CARNEGIE MUSEUM. looped. Epidermis brownish, usually painted with hair-like rays, forming here and there squarish spots. Hinge-teeth well developed. Male and female shell alike externally, but internally the female shell has an oblique depression for the marsupium. Soft parts with the inner lamina of the inner gills variable, free, except at the anterior end, to entirely connected, with all intergrades between these two extremes. Edge of the mantle not differentiated in front of branchial. Marsupium formed by the whole of the outer gills, with more crowded septa than the non-marsupial gills. Ovisacs only slightly extended beyond the edge of the gill, occupying only the marginal part of the gill, rather short, subcylindrical, and club-shaped (swollen at distal end); the whole marsupium is thrown into a number of folds (six to twenty). Placente very solid. Glochidia suboval, rather small. Type: P. phaseolus (Hildreth). This genus, in many respects, is the most primitive among the Lampsiline, but the folds of the marsupium represent a special structure. Ptychobranchus phaseolus (Hildreth). I have seen many specimens from the Ohio and Lake Erie drainages in Pennsylvania, and one gravid female from the Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler, coll. Febr. 6, 1911). Bradytictic, gravid from autumn to spring. The soft parts have been described and figured by Lea (Obs., VII, 1860, pl. 29, fig. 101) and Lefevre and Curtis (1910, pl. 1, fig. 1). Edge of mantle closed between the anal and supra-anal, the con- nection is short, but was never found missing. The branchial has papille, the anal is finely crenulated. In front of the branchial opening the inner edge of the mantle is first finely crenulated, but then becomes entirely smooth. Palpi of usual shape, their posterior margins con- nected for about one-fourth of their length. Gills long and moderately wide, the inner the wider. Their anterior attachment as usual, with the end of the inner gill slightly in advance of that of the outer, but widely separated from the palpi. Diaphragm normal, inner lamina of inner gill very variable: generally it is more or less free, and may be attached to the abdominal sac only at the anterior end, or for a greater distance; but in one case (out of thirty-two) it was found to be entirely connected. Thus, in this species, this char- acter is inconstant. ORTMANN: FAMILIES AND GENERA OF NAJADES. 307 Septa of the non-marsupial gills as usual. Marsupium formed by the outer gills in almost their whole length; only small sections are left free anteriorly and posteriorly; but in young individuals larger sections are non-marsupial. In the basal half the whole outer gill is non- marsupial, and has rather wide water-tubes, but the marginal half becomes marsupial, with much narrower water-tubes (ovisacs), and along the edge of the gill the ovisacs bulge out beyond it. This bulging out is only moderate. The placente are in the distal half of the gill; they are subcylindrical and club-shaped, being thicker toward the edge. The whole marsupium is thrown into a number of folds, increasing its surface, and further, in the distal part of the gill, the filaments are stretched or flattened out, so that the membranes en- closing the placenta become much thinner in this region than usual. Along the edge of the marsupium, the protruding ovisacs appear as a folded series of beads. The number of the ovisacs and of the folds is variable, and increases with age. Also in the sterile female the beads and folds are indicated on the edge of the marsupium. Placentz quite solid and permanent. They are discharged whole through holes formed at the end of the ovisacs (repeatedly observed). A brown stain is developed in the placentz, chiefly on their surface, which possibly indicates a hardening of the gelatinous matter. The eggs and glochidia are imbedded uniformly through the placental mass, but they are most crowded at the swollen ends. Glochidia (see Lea, Obs., VI, 1858, pl. 5, fig. 12;34 and Ortmann, 1911), pl. 89, fig. 14) rather small, subovate, without hooks, higher than long. Length 0.17; height 0.19 mm. Color of soft parts whitish, foot grayish, gills whitish, or grayish brown. Edge of mantle brown, broadly black posteriorly. Marsu- pium, when charged, blackish or purplish brown, inclining sometimes more to blackish, sometimes more to purple. Beads at edge more vividly colored, red or purple. A line of black markings near the edge, immediately below the protruding beads, on each side. 34 The poor quality of Lea’s figures of the glochidia is clearly shown in this in- stance. The figure of the glochidium of phaseolus (fig. 12) stands next to that of Eurynia recta (fig. 11), and is distinctly larger than the latter, while actually the glochidium of E. recta is by far the larger of the two. Also the outlines of these two glochidia are not quite correctly rendered. 308 ANNALS OF THE CARNEGIE MUSEUM. Ptychobranchus foremanianus (Lea). The soft parts of this species have been described by Lea (Obs., X, 1863, p. 443), and have been figured by him under the synonym woodwardianus (Obs., VII, pl. 29, fig. 103), and this species surely belongs in this genus. Ptychobranchus clintonensis Simpson. This species also belongs here, as is shown by Simpson’s description (19004, p. 79). Ptychobranchus subtentus (Say). Soft parts of one male and one gravid female were received from the Cumberland River, Burnside, Pulaski Co., Kentucky (B. Walker). Fic. 15. Ptychobranchus subtentus (Say). Gravid female from Cumberland River, Burnside, Pulaski Co., Ky. (Carn. Mus., No. 61, 4,971.) (Anal and supra- anal conjectural.) The soft parts of this species are entirely like those of P. phaseolus. The only difference I detect is in the extent of the marsupium, which consists of only five folds in my specimen, and does not occupy all of the outer gill, but leaves free a small section in front, and a somewhat larger one behind. Since I have only one female, I cannot tell whether this is always so, but I think it is unimportant, since likewise in P. phaseolus a considerable portion of the posterior end of the outer gill is non-marsupial in young individuals.*® The anal seems to be almost smooth, and nothing can be said about the supra-anal, since these parts are badly injured in both specimens. Posterior margins of palpi connected only at base. Inner lamina of inner gills in both % This is most noticeable in a young gravid female of P. phaseolus from Arka- delphia, Arkansas. ORTMANN: FAMILIES AND GENERA OF NAJADES. 309 specimens free, except in front, where it is connected for a short dis- tance in the female, and for a somewhat longer distance in the male. The glochidia are like those of P. phaseolus, but slightly larger. Length 0.18; height 0.22 mm. (see Plate XIX, fig. 5). The color of the soft parts is the same as in P. phaseolus. Mar- supium blackish purple, pale along the beaded edge, with black mark- ings like those in P. phaseolus. This species is placed by Simpson (19008, p. 591) in the letras dus, but he states that the soft parts were unknown to him. If we disregard the peculiar sculpture of the posterior slope of this shell, the structure of the hard parts is very similar to that of P. phaseolus. In some shells of P. swbtentus in the Carnegie Museum I have even seen the slight depression inside of the shell, which corresponds to the mar- supium.,*® Genus OBLIQUARIA Rafinesque. (1820.) (Simpson, 1900), p. 610.) Shell rounded oval, inflated. Disk with a row of large knobs, running from the beak to the center of the base, those of one valve alternating with the knobs of the other. Posterior slope corrugately sculptured. Beak-sculpture consisting of two or three rather heavy, but not sharply defined, concentric bars, which seem to be continued by the knobs of the disk. Epidermis greenish-yellow to brown, painted with numerous, delicate, wavy, and broken rays, which may be entirely absent. Male and female shells essentially alike. Inner lamina of inner gills free, except at the anterior end. Edge of mantle not differentiated in front of the branchial. Marsupium consisting of a few (generally less than ten) ovisacs, occupying a position just behind the center of the outer gill, beginning near the base of the gill, and reaching far beyond the edge. Thev are large, subcylindrical, and slightly curved, and have the rather solid placente of the same shape. Glochidia lying all through the placental mass, of medium size, almost subcircular. Type O. reflexa Rafinesque. Obliquaria is a primitive genus of the Lampsiline, which, in order to solve the problem of the aération of the glochidia, has reduced the 36 Sterki (1898, p. 31, and 1903, p. 103) describes in Ptychobranchus ‘‘a deep, oblique sulcus on the inside of each valve in the female, the space occupied by the marsupium.’’ This sulcus very often consists of a series of impressions correspond- ing to the marsupial folds. The same feature is observable in P. subtentus. 310 ANNALS OF THE CARNEGIE MUSEUM. marsupium to a few ovisacs, compensating for the reduction in their number, by an increase in their length. The tendency to move the marsupium backward toward the branchial opening is but slightly indicated. Obliquaria reflexa Rafinesque. Three males and one sterile female from the Ohio in Beaver Co., Pennsylvania, and another sterile female taken at Portsmouth, Scioto Co., Ohio, have been collected by myself. I received a sterile female from the Ouachita River, Arkadelphia, Clark Co., Arkansas, collected by H. E. Wheeler, February 6, 1911; and two males and four gravid females from Bayou Pierre, De Soto Parish, Louisiana, collected by L. S. Frierson, August 6, 1910. Soft parts described by Lea (Obs., X, 1863, p. 429) and figured by Lefevre and Curtis (1910, pl. 1, fig. 3). Branchial, anal, and supra-anal as usual, mantle connection be- tween the two latter moderately long, but shorter than the small anal. Branchial with papilla, anal crenulated. Toward the front the papilla of the branchial disappear suddenly, and the edge of the mantle is smooth, with a few indis- tinct crenulations just in front of the branchial. Palpi normal, their Fic. 16. Obliquaria reflexa Rafin- esque. Gravid female, from Bayou Pierre, De Soto Parish, La. (Carn. Mus., No. 61, 4,755.) at base. Gills short and broad, the inner posterior margins connected only wider anteriorly. Diaphragm normal; inner lamina of inner gills free from abdominal sac except at anterior end, where a portion less than half the length of the abdominal sac is connected. Anterior ends of gills normal. Septa and water-tubes of both gills of the usual structure in the male. In the female, the marsupium is formed by a part of the outer gill lying just behind the middle of the gill, and not extending to the posterior end of it. The most characteristic feature is that the mar- supium consists only of a small number (four to six in my specimens, but reported up to eight) of ovisacs, which, when empty, are hardly ORTMANN: FAMILIES AND GENERA OF NAJADES. 311 narrower than the normal water-tubes, but have much heavier septa, with the usual marsupial structure of the epithelium. When charged the ovisacs swell so as to be considerably wider than the normal water-tubes. The marsupium protrudes beyond the original edge of the gill to a considerable degree, and the single ovisacs are subcylin- drical, and are somewhat curved backward. The ova and glochidia fill the ovisacs in the shape of rather solid placenta, and are packed close together through all of the placental mass (see Lefevre.and Curtis. 1910, pl. 4, fig. 28). Probably the subcylindrical placente are dis- charged whole, although this has not been observed. The placentz can be taken out whole (see zbid., pl. 4, fig. 26), and in one of my specimens the holes through which placente had been recently dis- charged, have been seen (see Plate XVIII, fig. 10). The marsupium begins near the base of the gill, so that a considerable part of it is enclosed within the two original lamine of the gill. Lefevre and Curtis (1910, p. 97, fig. M) have figured the glochidium, and give its dimension as 0.225 X 0.23 mm. I find that this is sub- stantially correct. The glochidia are of medium size, almost sub- circular (their shape may be best compared with a circle a small section of which is cut off). Length and height about the same: 0.22 mm. (see Plate XX, fig. 1). Color of soft parts whitish, with the edge of the mantle brownish, chiefly so in the region of the branchial and anal openings. Mar- supium white. There is much uncertainty as to the breeding season of this species. Lea (Obs., II], 1842) mentions ova as formed in the ovariumin autumn, and Sterki (1898, p. 20) found them in October. Lefevre and Curtis (1910, p. 89) place this species among the forms with a short breeding season, but without giving particulars. The sterile females collected by myself were all found in the month of September, at a time when most other bradytictic forms are gravid. The gravid females from Louisiana, collected by Frierson on August 6, were in part discharging, so that this would tend to show that the breeding season ends at that time. The statements made by Lea and Sterki might suggest that the breeding season begins rather late, in winter, and this assumption would agree with the facts at hand. Yet recorded observations are entirely too few, and attention should be directed to this question. The sterile specimen from Arkansas, collected in February, does not centribute to the solution of the question. 33) WY ANNALS OF THE CARNEGIE MUSEUM. According to its known affinities and the gill-structure, this species should be bradytictic, and not tachytictic, as Lefevre and Curtis believe. But its primitive character makes it appear possible that in its breeding habits it may also be primitive, although I do not believe that it is a characteristic tachytictic form, for it possesses adaptations to a long breeding season. Genus CyPROGENIA Agassiz. (1852.) (Simpson, 1900), p. 609.) Shell rounded-triangular, inflated, often with a posterior ridge and a depression in front of it (especially in the young shell). Disk with nodular sculpture. Beak-sculpture obsolete (according to Simpson) slightly double-looped.” Epidermis greenish-yellow, painted with delicate rays, which break up into mottlings and spots. Male and female shell alike. Inner lamina of inner gills free from abdominal sac, except at anterior end. Edge of mantle in front of branchial with fine crenu- lations, which soon disappear anteriorly, but without special structures. Marsupium consisting of rather few (generally less than ten) ovisacs, lying in the center, or a little before the center, of the outer gills. The ovisacs begin near the base of the gill, and reach far beyond the edge. They are extremely long, and coil up spirally, in a backward and inward direction. The placente are very solid, subcylindrical like the ovisacs, and spiral. Glochidia distributed all through the placental mass, of medium size, almost semicircular. Type C. irrorata (Lea). The structure of this genus can easily be traced back to Obliquaria. The same general plan is observed in the structure of the soft parts, except that the marsupium is unusually elongated, and, in order to be accommodated in the shell, it is coiled up. Cyprogenia irrorata (Lea). I collected, September 24, 1910, two males and one gravid female in the Ohio River at Portsmouth, Scioto Co., Ohio, and received from B. Walker, three gravid females from the Cumberland River in Cumberland Co., Kentucky. No particulars as to the breeding season are known, but my specimen 87 Although I have several specimens with tolerably well preserved beaks, I have never seen the beak-sculpture clearly. ORTMANN: FAMILIES AND GENERA OF NAJADES. 313 from Portsmouth had eggs only, and thus the beginning of the breeding season is shown to be in autumn. The soft parts have been described and figured by Lea (Obs., I, 1834, pl. 5, figs. 6 and 7; and Obs., X, 1863, p. 433), but the figure is very poor.*® Branchial, anal, and supra- anal as usual, the latter two separated by a very short mantle-connection. Brarich- ial with papille, anal finely crenulated. In front of the branchial, the inner edge of the mantle has a series of fine crenulations which soon di. - appear, this edge becoming smooth. Palpi normal, pos- ieee A . P 1 | EiGes ize Cyprogenia irrorata (Lea). terior margins connected at Gravid female, from Ohio River, Ports- base only. mouth, Scioto Co., O. (Carn. Mus., No. Gills short and broad, the 61, 4,763.) inner much wider than the outer throughout its whole length. Diaphragm normal, inner lamina of inner gills free from abdominal sac, except at the anterior end. Anterior attachment of gills as usual. Septa and water-tubes in both gills normally developed, the latter moderately wide in the male and the non-marsupial gills of the female. Marsupium formed by a section in the middle of the outer gill; in fact this section is a little more toward the anterior end of the gill. Ovisacs few (three to eight in my specimens; up to eleven reported by other authors; Simpson gives for the genus twenty-three as maxi- mum), hardly different in width from the rest of the water-tubes, that is to say in the longitudinal direction. But, when charged, they swell somewhat in the transverse direction, so as to become subcylin- drical. The ovisacs project to an extreme degree beyond the edge of the gill. Although they begin near the base of the gill, and although a considerable part is enclosed between the original lamine of the gill, 38 Lea’s figures are quite characteristic of the marsupium itself, but the position of the latter in the animal (fig. 7) is wrong. Apparently the anterior and posterior ends of the body are inter-changed. The marsupium does not coil forward, as this figure shows, but backward. 314 . ANNALS OF THE CARNEGIE MUSEUM. this part is very small when compared with the prolonged portion. The latter curves backward in a circle, and is rolled up spirally, the spiral forming about one and a half to two turns, but only the posterior ovisacs complete the whole revolution, while the anterior ones stop earlier, the first after completing the circle about once. The distal parts of the spiral wind up in the direction toward the median line of the body, so that in a view from the outside, they are hidden under the outer gill and the first whorl of the marsupium. The ova fill the ovisacs in the shape of closely packed masses, forming distinct and very solid placentae, red in color, rarely white. Glochidia rather small, almost semicircular, distinctly longer than high, without hooks. Length 0.18; height 0.15 mm. (see Plate XIX, fig. 6). Sterki (1898, p. 19) gives the dimensions as length 0.21; height 0.17; diameter 0.14 mm. He also says that the glochidial shell is ‘considerably longer than high and has numerous distinct, crowded, concentric lines of growth.”’ I have not seen the latter. The shape of the glochidia approaches to a degree, that of Dromus, but the disproportion between length and height is much less. Color of soft parts whitish. Abdominal sac and mantle suffused with black. Edge of mantle brown with black spots, this mottling extending all around. Marsupium, when charged, red, or (according to Sterki) sometimes white. Genus Dromus Simpson. (1900.) (Simpson, 1900), p. 614.) Shell very much like that of Cyprogenia. Beak-sculpture obsolete, described by Simpson as consisting of interrupted, concentric ridges, but I have never seen them distinctly. Inner lamina of inner gills partly free from abdominal sac, connected near the anterior end for about one-third, or more, of the length of the abdominal sac. Edge of the mantle in front of branchial without special structures. Marsupium consisting of numerous ovisacs, which occupy the larger posterior portion of the outer gill, leaving a smaller anterior section non-marsupial. The ovisacs are comparatively short, subcylindrical, or only slightly compressed, and lie practically entirely beyond the original edge of the gill. In older individuals, the mar- supium becomes warped and folded. Placentz solid, subcylindrical, or slightly compressed, rather short. Glochidia placed chiefly toward ORTMANN: FAMILIES AND GENERA OF NAJADES. B35) the outer walls of the marsupium, around the edges of the placenta, which are central. Glochidia of peculiar shape, small, much longer than high. Type D. dromas (Lea). This is a highly interesting genus, = several quite unique features (general shape of marsupium, arrangement of glochidia and placente, shape of glochidia), bu1 there is no question that it belongs to the more primitive types of the Lampsiline. In young specimens, where the marsupium is not folded, the marsupium resembles somewhat that of Obovaria, but without being so swollen. The general shape of the shell, as well as the shape of the glochidia can only be compared with that of Cyprogenia. I do not think that the folded marsupium in- dicates closer relationship to Ptychobranchus, since here the folds are of a different character. In the absence of special structures on the edge of the mantle, this genus shows only a low stage of specialization, without pointing to any particular affinity with other forms. Dromus dromas (Lea). I am indebted to B. Walker for seven complete specimens, and the soft parts of nine others, all from the Cumberland River, in Pulaski, Russell, Wayne, and Cumberland Cos., Kentucky. One of the soft parts was a male, the others were all gravid females with glochidia. They all were collected late in the season of 1910, so that the beginning of the breeding season is in autumn. The description of the marsu- pium given by Simpson (1900), p. 615) is entirely inadequate and directly misleading, and in one particular (‘‘bases of the ovisacs slightly rounded’’) un- intelligible HRKe “antes Dromus dromas (Lea). Awal o araved oft Gravid female, young (shell 41 mm. Be OPES HP < om long), from Cumberland River, Albany the supra-anal by a very short Landing, Cumberland Co., Ky. (Carn. and deciduous mantle-connec- Mus., No. 61, 4,970.) tion: only in two young and one older specimen was the latter preserved; but the others had been rather roughly handled and the posterior region of the mantle was 316 ANNALS OF THE CARNEGIE MUSEUM. generally more or less injured.*® Anal large, with crenulations. Branchial large, with papillae; toward the front papillae gradually changing to crenulations, which soon disappear, so that the anterior inner edge of the mantle is smooth. Palpi of usual shape, but small; their posterior margins united for about one-half, or less, of their length. Gills conforming to the shape of the shell, rather short and broad, the inner much the wider anteriorly. The anterior attachment of the gills is as usual. Gill-diaphragm normal. Inner lamina of inner gills free from abdominal sac, except anteriorly, where it is connected for about one-third to almost one-half of the length of the abdominal sac. Fic. 18a. Gravid female, medium size (shell 55 mm. long), from Cumberland River, Eadsville, Wayne Co., Ky. (Carn. Mus., No. 61, 4,968.) Gills with well developed water-tubes and septa. Those of the male, and the non-marsupial gills of the female, with distant septa and wide water-tubes. Marsupium formed by the posterior section of the outer gill; more than half of the gill takes part in it; a larger section in front and a smaller behind remain non-marsupial. The marsupial part bulges out considerably beyond the original edge of the gill, about as wide again as the gill, and in this section the septa are much more crowded, and the ovisacs are narrow. When gravid, the ovisacs swell only slightly, so that they are very little compressed, and chiefly so near their base. The placente have the same subcylindrical and only slightly compressed shape. Practically the whole of each ovisac 39 But in some it was positively absent. This is also a rather primitive condition, not observed in any*other form of the Lampsiline. ORTMANN: FAMILIES AND GENERA OF NAJADES. 317 lies in the outbulging part of the gill, and only the basal ends extend very little in between the original gill-lJamine.” In young specimens, the marsupium is simple, and forms a smooth, compressed (not much swollen) body, marked off from the anterior and posterior non-marsupial parts of the gill by irregular folds. In larger specimens, however, this marsupial mass begins to warp, and finally is folded up into a number of irregular folds. The strongest folds are near the anterior end of the marsupium. In none of my specimens does the marsupium occupy the whole margin of the gill. The placente are quite solid and permanent, and possess a_ peculiar structure (see Plate XVIII, fig. 11). In all specimens at hand glochidia are developed, and they appear arranged around a central axis (placenta), the color of which is white or red. The latter color, if present, is restricted to this axis, and the glochidia themselves are transparent white, and form a fringe around the narrow edges of the placente. They seem to be connected with them by fine threads, possibly Fic. 18). Left gills of a large gravid female (shell 60 mm. long), from Cumberland River, near Ro- wena, Russell Co., Ky. (Carn. their embryonal threads.“ Since the Mus., No. 61, 4,966.) placentze themselves almost touch the septa, the glochidia are thus crowded toward the lateral faces of the marsupium. Whether this arrangement is already present in the eggs, is unknown to me. It is, however, certain that this arrangement can only have the purpose of bringing the glochidia as close as pos- sible to the wall of the marsupium, in order to give them the best chance to be near the current of fresh water going over the mar- supium. This is one of the little special devices for the proper aération of the glochidia. Glochidia of unique shape; they are much longer than high, and 40 Simpson says that the marsupium occupies the “‘ base”’ of the outer gills. This is a very ambiguous expression, but apparently is intended to imply that it is situated on the margin of the gills, while the ‘“‘base’’ is non-marsupial. This is a very peculiar feature of Dromus, and not met with in any other genus. Only Ptychobranchus has the same condition slightly indicated. 41 The glochidia adhere rather firmly to the placental mass by their threads and it is hard to isolate them, except with caustic potash. 318 ANNALS OF THE CARNEGIE MUSEUM. might be called bean-shaped. No hooks are present. Length 0.19; height 0.10 mm. (see Plate XIX, fig. 7). Color of soft parts whitish. Foot yellowish white, basal part (abdominal sac) gray or blackish. Gills gray or grayish white. In the gravid female, the marsupium is white or red. Mantle more or less suffused with black, whitish toward margins and front parts. Its edge has alternating chestnut-brown and black spots. Anal opening inside of this maculated edge with a white, followed by a black band. Genus FRIERSONIA gen. nov. Shell subelliptical, without distinct posterior ridge. Disk not sculptured. Beak-sculpture of the double-looped pattern, consisting of six to eight fine bars, of which the later ones are distinctly double- looped, and the latest are interrupted (unconnected) in the middle. Epidermis greenish-yellow, with rather distinct, simple rays. Male and female shells hardly different. Inner lamina of inner gills connected with abdominal sac. Edge of mantle in front of branchial slightly lamellate, with fine and distinct crenulations, disappearing gradually in front, but without papille. A brown streak of pigment along this part of the edge. Marsupium consisting of many ovisacs, occuping the larger posterior section of the outer gill. When gravid, the ovisacs swell very little, and they are only slightly compressed in the basal part, which is largely enclosed between the lamine of the gill. The ovisacs reach considerably beyond the edge of the gill, and in this region they are curved backward in a peculiar manner, subcylindrical, and tapering toward a point directed backward at the hind end of the marsupium. The marsupium has also a remarkably sharp edge. Placentz not very solid. Glochidia lying all through the placental mass, of medium size, and subovate in shape. Type F. iridella (Pilsbry and Frierson). According to the arrangement in the key (p. 304) this genus would appear to fall into the same group with the preceding genera. But this is hardly the case. It has in common with the genera with which it has been associated in the key only the fact that the marsupium is not of the simple kidney-shape shown by the genera which follow in the key. The sharp edge of the marsupium, its posterior point, and the recurved ovisacs are quite unique. For the present, I do not under- stand the meaning ot this structure, but it may be connected with the ORTMANN: FAMILIES AND GENERA OF NAJADES. 319 discharge of the glochidia. In shell characters Friersonia shows nothing very characteristic, but it approaches more nearly the Nephronajas- and Eurynia-types than any other. Friersonia iridella (Pilsbry and Frierson). Lampsilis iridella, Pilsbry and Frierson, 1908, p. 81 (figure published in 1907, on pl..12); Pilsbry, 1909, p. 534. I have the soft parts of three gravid females, one with eggs, two with glochidia, from Valles River, Valles, San Luis Potosi, Mexico, and owe them to the courtesy of L. S. Frierson. They were collected by A. A. Hinkley in December and January, 1906-1907.% Cotypes of species. Fic. 19. Friersonia iridella (Pilsbry and Frierson). Gravid female, from Valles River, Valles, San Luis Potosi, Mexico. (Carn. Mus., No. 61, 4,495.) To the characters of the soft parts mentioned in the generic diagnosis the following should be added: Anal and supra-anal separated by a rather short mantle-connection (these parts are greatly injured in all three specimens, but the remnants of the connection may be seen in one specimen). Anal finely crenulated. Branchial with papille. Palpi rather large, of usual shape, posterior margins connected about one-half of their length. Diaphragm and anterior attachment of gills normal. Gills of gravid female rather long, the outer marsupial gill covering all of the inner gill, except its anterior end. Septa and water-tubes as usual. Marsupium quite long, and formed by a greater part of the outer gill than usual, yet there is a portion at the anterior end and a very small one at the posterior end, which are non- marsupial. Ovisacs over fifty. The sharp edge of the marsupium 42 See Hinkley, 1907, p. 68. 320 ANNALS OF THE CARNEGIE MUSEUM. and the peculiar posterior point are quite evident in all three specimens, and the same is true of the peculiar curve of the ovisacs, so that these features cannot be accidental. Glochidia higher than long. Length 0.20; height 0.22 mm. (see Plate XIX, fig. 8). Colors largely faded in my alcoholic material. One specimen has a peculiar brown line across the middle of the foot. The edge of the marsupium has brownish black pigment in spots. This species and genus is one of the most peculiar with which I am acquainted. I would have considered it a Nephronajas, but since I have investigated another Mexican Nephronajas (see below), it is clear it cannot belong to this genus. The generic name is selected in recognition of the valuable help in my work received from Mr. L. S. Frierson. Genus OBOVARIA Rafinesque. (1819.) (Simpson, 1900), p. 599.) Shell rounded or ovate, higher than long, or only slightly longer than high, inflated, without distinct posterior ridge. Disk not sculp- tured. Beak-sculpture poorly developed, consisting of few subcon- centric bars, of which the later ones have sometimes the tendency to become sinuate, but are not distinctly double-looped. Epidermis yellowish to brownish, rarely greenish, with indistinct, simple rays or without rays. Male and female shell slightly different in shape, the female being generally a little expanded on the post-base, but this difference is sometimes hardly noticeable. Inner lamina of inner gills entirely connected with abdominal sac. Edge of the mantle very little differentiated in front of the branchial. It is slightly lamellar, with fine crenulations; and this part is generally emphasized only by the thickening of the margin of the mantle and the presence of a streak of dark pigment; there are never papille on it. Marsupium consisting oi many ovisacs, occupying the posterior part of the outer gill. The ovisacs, when charged, swell transversely, so as to become lanceolate and compressed. They reach from near the base of the gill to, and a good deal beyond, the edge of the gill, and the whole marsupium assumes a kidney-shape. Placentz not very solid. Glo- chidia all through the placental mass, of medium size and subovate. Type O. retusa (Lamarck). This is another primitive type of the Lampsiline, leading, however, toward the more highly developed forms of the subfamily. The ORTMANN: FAMILIES AND GENERA OF NAJADES. 321 marsupium in this genus is not very peculiar, though assuming the characteristic kidney-shape of the higher Lampsiling. The task of aérating the glochidia is taken up by the edge of the mantle in front of the branchial opening. However, the latter is as yet very little differentiated morphologically, but the thickened (muscular) margin and the pigment indicate that it actually has a special function. The shell of Obovaria presents no remarkable features, though it is possibly archaic, for it reminds of certain forms of Fusconaja and Quadrula. This genus is divided by Simpson into two subgenera, which are very well defined. Subgenus Obovaria (sens. strict.). Shell rounded, rather upright, beaks more or less in the middle of the upper margin. Pseudocardinals normal and divergent. Type O. retusa (Lamarck). Subgenus Pseudodn Simpson (1900), p. 601). Shell ovate, oblique, beaks quite anterior. Pseudocardinals (at least in old shells) oblique, almost parallel to the laterals. Type O. ellipsis (Lea). At first glance, O. ellipsis looks very different from typical Obovaria, and I was for some time inclined to unite it with Nephronajas; but O. castanea clearly forms a connection with the typical forms. Simpson (/. c.) describes the soft parts of Pseudoén, and says: “mantle having a wide, thickened, double border, the inner edge being toothed throughout below.’’ This is incorrect. The inner edge is slightly widened and crenulated only for a short distance in front of the branchial: Simpson also says that the ovisacs are ‘‘tinted with purple below.”’ I have not seen this in O. castanea, although O. ellipsis hasa slight purplish gray pigment at the edge of the marsupium:; but this should not be described as “‘ purple.” Obovaria retusa (Lamarck). On August 29, 1908, I found a gravid female with eggs in the Ohio River in Beaver Co., Pennsylvania, and on September 22, 1910, I secured two males and two gravid females, with glochidia, in the Ohio River at Portland, Meigs Co., Ohio. The soft parts have been described by Lea (Obs., X, 1863, p. 433). Anal and supra-anal separated by a short mantJe-connection. Anal crenulated, branchial with papilla. In front of the branchial the inner edge of the mantle in the female is slightly dilated and lamelliform, oon ANNALS OF THE CARNEGIE MUSEUM. with fine crenulations. It is defined on the inner side by a narrow stripe of black pigment. In the male this lamella is also present, but much weaker. Farther in front the edge of the mantle is smooth. Palpi small, normal, their posterior margins connected only at the base. Gills short and broad, the inner ones broader. Diaphragm normal. Inner lamina of inner gills entirely connected with abdominal sac. Anterior attachment of gills as usual. Septa and water-tubes in both gills normally developed. Mar- supium restricted to a small section in the posterior half of the outer gill, leaving more than half of the anterior portion and a small posterior section non-marsupial. Ovisacs fifteeen to twenty (in my specimens); when charged not narrower than the normal water-tubes in the longi- tudinal direction, but expanding in the transverse direction, so that their lumen becomes lanceolate and compressed, the whole mar- supium thus appearing swollen and kidney-shaped. The marsupium extends considerably beyond the original edge of the gill, and about three-fourths of the length of the ovisacs is within the lamine of the Fic. 20. Obovaria retusa (Lamarck). Gravid female, from Ohio River, Port- land, Meigs Co., O. (Carn. Mus., No. 61, 4,773.) gill, while one-fourth lies beyond the latter. Edge of marsupium blunt, without pigment. Placente not well developed; eggs and glochidia rather loose. Glochidia rather large, suboval, without hooks. Length 0.22; height 0.27 mm. (see Plate XIX, fig. 9). Color of soft parts whitish, only edge of mantle brown, with a black streak in front of the branchial. Charged marsupium whitish. ORTMANN: FAMILIES AND GENERA OF NAJADES. Se} Obovaria circulus (Lea).* About a dozen specimens from the Ohio drainage of western Penn- sylvania have been investigated, among them gravid females. Several additional specimens were examined from the Ohio River in Ohio. Gravid females were found in the month of September, and on May 27, In the latter case, discharging individuals were secured, with holes in the edge of the marsupium. Structure of soft parts essentially the same as in O. retusa. In the sterile female, the ovisacs are slightly narrower than the regular water- tubes. Number of ovisacs up to thirty and more. As in the pre- ceding species the edge of the mantle in front of the branchial is in the female slightly lamellar and crenulated, but has only a brown (not blackish) mark along it. Glochidia (see Ortmann, 19110, pl. 89, fig. 15) similar to those of O. retusa, but smaller. Length 0.20; height 0.23 mm. Obovaria unicolor (Lea). I have received from A. A. Hinkley three males and eight gravid females, taken from the Pearl River, Jackson, Hinds Co., Mississippi, on Nov. 5, 1910. This species agrees in every particular with O. cir- culus, except that the glochidia are smaller. Length 0.16; height 0.21 mm. (see Plate XIX, fig. 10). Obovaria (Pseudo6n) ellipsis (Lea). Two females were collected in the Ohio River in Beaver Co., Penn- sylvania, on August 29, 1908, one sterile, the other just beginning to fill the marsupium with eggs. Three males, one sterile, and six gravid females were secured in September, 1910, in the Ohio River in Ohio. Thus the beginning of the breeding season is normal. The soft parts are much like those of the other species of the genus. Anal and supra-anal separated for a short diatance. Anal crenulated, branchial with papilla. In front of the branchial the inner edge of the mantle of the female is slightly lamelliform, with fine crenulations. This part does not reach to’the middle of the lower margin, and farther in front the edge is smooth. Posterior margins of palpi connected only at base. Inner lamina of inner gills connected with abdominal sac. 43 This includes O. lens (Lea), which is not specifically distinct. 324 ANNALS OF THE CARNEGIE MUSEUM. Marsupium formed by about the posterior half of the outer gill, kidney-shaped, consisting of as many as forty and more ovisacs, its edge slightly pigmented. Glochidia similar to those of the other species. Length 0.19; height 0.22 mm. (see Plate XIX, fig. 11). Color whitish, edge of mantle inclining to blackish, chiefly in the region of the branchial and anal, and more intense in the male sex. Pigment on edge of marsupium purplish gray, not sharply marked. Obovaria (Pseudoén) castanea (Lea). Twelve males, one sterile, and five gravid females (with glochidia) from the Ouachita River, Arkadelphia, Clark Co., Arkansas, have been sent by H. E. Wheeler. They were collected on February 6 and March 215 T9Rx. Identical in all essential respects with O. ellipsis. Marsupium formed by twenty to thirty ovisacs and its edge not pigmented. A grayish streak along the inner edge of the mantle in front of the branchial. Glochidia of the same shape as those of O. ellipsis, but smaller. Length 0.15 mm.; height 0.19 mm. Genus NEPHRONAJAS Crosse and Fischer. (1893.) (Simpson, 1900), p. 591.) Shell ovate or subelliptical, distinctly longer than high, compressed or slightly inflated, without, or with, indistinct posterior ridge. Disk not sculptured. Beaks moderately anterior, never in the middle of the shell, and never very near the anterior end. Beak-sculpture poorly developed, consisting of a few faint bars, which have a tendency to become double-looped, with the central part between the loops obliterated. Epidermis yellowish to greenish, generally with distinct 44 There is some doubt as to the identity of my specimens. B. Walker has a number of sets of a shell from Alabama, Mississippi, Louisiana, and Arkansas, of which he sent me specimens, and some of which have been labeled by Simpson castanea, but which are certainly different from the present form, and probably do not belong to this genusat all. Although I have not seen Lea’s type, I believe that I have the real castanea, for the reason that all authors (Lea, Obs., I, 1834, p. 91; Call, 1895, p. 9; Simpson, 1900), p. 602) who have discussed this species, emphasize its similarity to O. ellipsis. Lea’s words: “‘ This small species is allied to U. circulus (nob.) in colour and to U. ellipsis (nob.) in form”’ are entirely sufficient to recognize it. There is no other form known to me, of which this could be said. Also Vanatta (1910, pp. 102 and 103) quotes O. castanea from the Ouachita River in Arkansas, ORTMANN: FAMILIES AND GENERA OF NAJADES. 325 green rays. Male and female shells differing in shape, but the dif- ference often hardly noticeable. Soft parts agreeing with those of Obovaria in every respect; the glochidia also of the same type. Type JN. plicatula Charpentier. In its anatomical structure this genus is indistinguishable from Obovaria. The differences are all in the shell. But while Obovaria is primitive in its shell, Nephronajas inclines toward the genera of the Lampsilis-type, in fact its species have been considered, at least temporarily, to belong to Lampsilis. The subgenus Pseudoén of Obovaria connects this genus with Obovaria, and so we have an almost complete series. The nomenclature of this genus is doubtful. Of the species, which belong here, two (/igamentina and perdix) stand according to Simpson's system in Lampsilis, and a third (sapotalensis) in Nephronajas. Since Lampsilis is retained for other forms, only Nephronajas is available. But the anatomy of the type species of Nephronajas (plicatula) remains as yet unknown, and it is possible that it may differ in anatomy from sapotalensis. In the latter case, of course, Nephronajas could not be used for the present genus, and a new name would have to be chosen. Nephronajas ligamentina (Lamarck). Numerous specimens from the Ohio drainage in western Penn- sylvania have been investigated. In addition specimens have been seen from the Ohio River in West Virginia and Ohio (collected by myself), from the Cumberland River in Kentucky (var. gibba), received from B. Walker, and from the Ouachita in Arkansas, received from H. E. Wheeler. Typically bradytictic. The breeding season begins in August, and specimens with eggs are present in this month and the beginning of September. Later on only glochidia are observed. Specimens with glochidia have again been observed in May, and the discharge must take place in this month, for in June and July no gravid females have ever been found, although numerous specimens have been in- vestigated. .The soft parts have been discussed by Lea (Obs., X, 1863, p. 424) and Simpson (in Baker, 1898, p. 108). Anatomy in every particular like that of Obovaria. The inner edge 326 ANNALS OF THE CARNEGIE MUSEUM. of the mantle of the female in front of the branchial is very slightly lamellate and finely crenulated. It is even less developed than is generally the case in Obovaria. The edge of the mantle is brown all around, often very pale, often darker, and becomes blackish in the region of the branchial and anal. Marsupium generally quite large and swollen, with as many as forty ovisacs, or even more. At its edge there is generally brownish or blackish pigment, but this may be indistinct, or even lacking. For an account of the glochidia see Lea, Obs., VI, 1858, pl. 5, fig. 18; Ortmann, 19110, pl. 89, fig. 16. Length 0.22; height 0.24. Lefevre and Curtis (1910, pl. 4, figs. 24 and 27) have figured the placenta, but they are distinguishable only when the eggs are present, later the cohesion is lost. Nephronajas perdix (Lea). I have received three gravid females from B. Walker. They are from the Cumberland River, Burnside, Pulaski Co., Kentucky. This species agrees completely with NN. ligamentina, and with Obovaria in general. _ In this species also the inner edge of the mantle of the female in front of the branchial is slightly lamellar and indis- tinctly crenulated, and emphasized by a streak of black pigment. Glochidia rather large. Length 0.25; height 0.29 mm. (see Plate XG hig. 2): The affinity of this species with JN. ligamentina has been recognized by Lea and Simpson, and thus it is not astonishing that the anatomy should prove to be the same. Nephronajas sapotalensis (Lea).” Three males, and two sterile females, from Hueyapam River, Hacienda de Cuatotalapam, Canton Alayucan, State of Vera Cruz, Mexico, taken July 23, 1910, have been examined. I received these specimens from A. G. Ruthven, and they belong to the Museum of the University of Michigan at Ann Arbor. One female has been kindly deposited in the Carnegie Museum. 4 B. Walker writes to me about these specimens: they ‘‘agree exactly with Crosse and Fischer’s figure of their computatus, which according to von Martens is probably only a variety of sapotalensis, differing mainly in having the pseudocardinals slender, while in sapotalensis they are heavy.’’ The type locality of sapotalensis is Sapotal River, near Tlocatalpam, Mexico. This is not far from the locality of my speci. mens, and in the same general drainage system (Papaloapan and San Juan Rivers) - ORTMANN: FAMILIES AND GENERA OF NAJADES. 327 In every respect like the two preceding species. Mantle-connection between anal and supra-anal shorter than anal, the latter with crenu- lations, the branchial with papilla. Inner mantle edge of the female in front of branchial very slightly lamellate, with fine crenulations, and a black streak accompanying it. Palpi with posterior margins con- nected at base. Inner lamina of inner gills entirely connected with abdominal sac. Marsupium formed by numerous (twenty and more) water-tubes, occupying a section of the posterior half of the outer gill. In the sterile females at hand, the margin of the mar- supium is rounded, projecting slightly beyond the edge of the gill, beaded, and marked with brownish black pigment. Placente and glochidia unknown. Fic. 21. Nephronajas sapotalensis (Lea). Sterile female, from Hueyapam River, State of Vera Cruz, Mexico. (Carn. Mus., No. 61, 5,000.) Color whitish, with the margin of the mantle brown and blackish in the region of the anal and branchial, a blackish streak in front of the branchial, and black pigment on the edge of the marsupium. Genus AMYGDALONAJAS Crosse and Fischer. (1893.) Simpson, 1900), p. 604 (as subgenus). Shell ovate-triangular, inflated, truncated at posterior slope, with a distinct and often sharp posterior ridge. Disk not sculptured. Beak-sculpture consisting of a few fine ridges, of which the later ones are more or less distinctly sinuated or double-looped. Epidermis yel- lowish green, with a pattern of broken or arrow-marked rays. Male and female shells differing but little, the female shell somewhat inflated. at the post-basal region. Inner lamina of inner gills entirely connected, or free for a short distance. Inner edge of mantle in front of branchial in the female, 328 ANNALS OF THE CARNEGIE MUSEUM. slightly lamellar for a certain distance, with fine crenulations. Mar- supium apparently like that of Obovaria (I have seen only sterile females). Glochidia (according to Lefevre and Curtis, 1910, p. 97, fig. G), of suboval outline, but extremely small. Type A. cognata (Lea), a Mexican species, of which the soft parts are unknown. This genus stands close to Obovaria and Nephronajas, and has essentially the same structure of the soft parts. It differs, however, in the shape of the shell, and most emphatically in the glochidia. A final definition of the genus depends on the investigation of the anatomy of the type-species. Amygdalonajas elegans (Lea). Two males were found in the Ohio River in Beaver Co., Pennsyl- vania, by myself. From L. S. Frierson, I received three males and three sterile females from Bayou Pierre, De Soto Parish, Louisiana, collected Aug. 6, 1910. The soft parts are described by Simpson (in Baker, 1898, p. 93). Anal and supra-anal separated by a rather long mantle-connection, about as long as the anal. Anal crenulated, branchial with papille. In front of branchial, the inner edge of the mantle in the female is narrowly lamellar, with fine crenulations, this part reaching to about the middle of the lower margin. Posterior margins of palpi connected for about one-third of their length. Gills and diaphragm of the usual shape. Inner lamina of inner gills sometimes entirely connected with the abdominal sac, sometimes free for a short distance (maximum about one-fourth the length of ab- dominal sac); often only small holes at posterior end of foot are left open. Gills of the usual structure. In the female the marsupium is formed by the posterior part of the outer gills (a little over one-half). Ovisacs numerous, projecting beyond edge of gill. Charged marsupium not observed. The glochidia have been described and figured by Lefevre and Curtis (1910, p. 97, fig. G), and are characterized by their extremely small size. Length 0.075; height,0.09 mm. Color of soft parts whitish, with the edge of the mantle blackish, mottled with black and white in the region of the branchial and anal. Along the inner edge in front of the branchial is a streak of black pigment. ORTMANN: FAMILIES AND GENERA OF NAJADES. 329 Genus PLAGIOLA Rafinesque. (1819.) (Simpson, 1900), p. 603.) Shell subtriangular, somewhat inflated, but peculiarly compressed toward the beaks, with a distinct posterior ridge, and a narrow, trun- cated, posterior slope. Disk not sculptured. Beak-sculpture indis- tinct, consisting of a few, fine, concentric, and slightly and indistinctly double-looped ridges. Epidermis yellow, greenish, or brownish, painted with rays, which are broken into lunate, or squarish, blotches. Male and female shells slightly different in shape, the female smaller, more inflated, and slightly swollen in the post-basal region. Inner lamina of inner gills free for a greater or smaller distance. Inner edge of mantle in front of branchial for a certain distance slightly lamellar and with fine teeth, but without papille in the female. Mar- supium like that of Obovaria, kidney-shaped. Placentz lanceolate, not very solid. Glochidia very large, spatulate, gaping at the anterior and posterior margins. Type P. securis (Lea). In the soft parts, this genus stands essentially upon the same stage of development as Obovaria and Amygdalonajas. Its chief character- istics are the shape of the shell and of the glochidia. The latter are quite unique, and possibly indicate a transition toward the glochidia of Proptera. Plagiola securis (Lea). About half a dozen specimens from the Ohio and Allegheny in western Pennsylvania have been collected by myself. I received from B. Walker a gravid female from the Cumberland River in Kentucky, and another from H. E. Wheeler from the Ouachita River in Arkansas. In September and October this species is regularly found gravid, so that the beginning of the breeding season is normal. The soft parts have been described by Lea (Obs., X, 1863, p. 43). Anal and supra-anal separated by a manile-connection of medium length. Inner edge of anal crenulated, that of branchial with papille. In the female, the inner edge of the mantle in front of the branchial is slightly lamellar, with fine teeth, which are rather distant, but with- out papillae. Posterior margins of palpi connected for about one- fourth of their length. Gills and diaphragm of usual shape. Inner lamina of inner gills more or less free. The maximum observed was free for about three- 330 ANNALS OF THE CARNEGIE MUSEUM. fourths of the length of abdominal sac, the minimum was only a small hole at the posterior end of the foot, but this was only on one side of the body; on the other side the lamina was free for a little less than half the length of the abdominal sac. Gills of the usual structure. Marsupium kidney-shaped when charged, occupying the posterior half or more of the outer gill, with numerous (thirty and more) ovisacs, of the Obovaria-type. Ovisacs compressed, lanceolate, with poorly developed placente. Glochidia distributed all through the placentae, of unusual shape (see Lea, Obs., VI, 1858, pl. 5, fig. 6;46 Lefevre and Curtis, 1910, p. 97, fig. H; Ort- mann, 19110, pl. 89, fig. 17). They are quite large, subspatulate, (dilated and rounded off toward the ventral margin), and their an- terior and posterior margins are distinctly gaping. Lefevre and Curtis give the measurements as follows: length 0.23; height.c.31; while my maximum measurements are: length 0.26; height 0.35 mm. Color whitish. Edge of mantle brownish black, chiefly so in the posterior region. The black pigment is emphasized along the edge in front of the branchial. Genus PARAPTERA Ortmann. (1911.) (Ortmann, 1911), pp. 301, 334, 338.) Shell thin, elliptical, or obovate, when young with a distinct pos- terior wing, rather compressed, without posterior ridge. Disk without sculpture. Hinge-teeth feebly and often imperfectly developed. Beak-sculpture fine, consisting of a few concentric bars, followed by a few others, which are double-looped. In the latter, only the posterior loop is distinct, while the anterior is obliterated. Male and female shells slightly different, the female shell more expanded at post-base. Inner lamina of inner gills entirely connected with abdominal sac. Edge of the mantle of the female slightly lamellar in front of branchial, with crenulations, but not with papillae. Marsupium kidney-shaped, swollen, consisting of many ovisacs occupying the posterior part of the outer gills. Placente not very solid. Glochidia very small, of suboval shape. Type P. gracilis (Barnes). Another genus having the structure of Obovaria, distinguished only by the shape of the shell and the glochidia. The latter are very remark- able, and can only be compared with those of Amygdalonajas. 46 This figure is not drawn to scale, and is much too small. ORTMANN: FAMILIES AND GENERA OF NAJADES., 331 Paraptera gracilis (Barnes).” I myself collected about a dozen specimens with soft parts in the Ohio and Lake Erie in western Pennsylvania, and received, from R. L. Moodie, a male from the Kansas River in Kansas, and from L. S. Frierson a male and a sterile female from Bayou Pierre, Louisiana. The breeding season seems to have certain peculiarities. The species is undoubtedly bradytictic, but the season begins rather late. I found specimens with the marsupium partly charged as early as August 30, but these, as well as others found in September and October, all had only eggs, but no glochidia. In spring, a discharging female has been observed as early as May 22, but others were found fully charged with glochidia as late as July 7, 8,and 11.8 Thus the breeding season must last from the end of August to about the middle of July, with only a very short interval. Incomplete descriptions of the soft parts have been published by Lea (Obs., X, 1863, p. 434) and Simpson (in Baker, 1898, p. 99). Anal and supra-anal separated by a mantle-connection which is slightly longer than the anal. Anal crenulated, branchial with papille. In the female the inner edge of the mantle in front of the branchial is lamellar, somewhat dilated, with fine crenulations, running forward for about one-third the length of the margin of the mantle. No papillae are present. Palpi with the posterior margins united at the base only. Gills and diaphragm of normal shape. Inner lamina of inner gills connected throughout with abdominal sac. Structure of gills normal. Marsupium formed by the posterior part of the outer gill, kidney- shaped, and swollen. Ovisacs numerous (thirty to forty), lanceolate. Placentz not very solid, glochidia distributed all through the mass, very small, of suboval shape. Length 0.08; height 0.09 mm. (See 47 Lea (Obs., VIII, 1862, p. 79, pl. 9, fig. 224) described U. dolosus, which Simpson (1900), p. 568) makesa synonym of U. purpuratus Lea, but states that the glochidia are pouch-shaped (Lampsilis-type), not wedge-shaped as in Paraptera purpurata. Simpson dismisses this by saying (J. c., footnote 2) that the ‘‘form of embryos ina given species is often not constant.’’ This is a very serious mistake, for there is nothing more constant for the species than the glochidium. Call (1895, p. 19) makes dolosus a synonym of gracilis, and I think that he is right. Simpson (I. c., Pp. 574) does not report gracilis from the Alabama drainage, but it surely is there (as dolosus). Ihave myself seen two specimens from the Coosa River at Wetumpka, collected by H. H. Smith in October, 1901. 48 Those collected July 7 and 8 are from Lake Erie, but the one collected July 11 is from the Ohio River. oon ANNALS OF THE CARNEGIE MUSEUM. Lefevre and Curtis, 1910, p. 97, fig. K, where the measurements given are: length 0.07; height 0.09; Ortmann, 19114, pl. 89, fig. 19, and Coker and Surber, 1911, pl. 1, fig. 2). Soft parts whitish. Edge of mantle in the region of anal and branchial brownish black. * 4 { 5 , . + . f - od a " ‘ ~ v > a) 7 > “A ‘ . TPA . i ae Pik “NAY ADES (FRESHWATER MUSSELS) OF “THE UPPER TENNESSEE DRAINAGE. WITH "NOTES ON-SYNONYMY AND DISTRIBUTION. By A. E. ORTMANN, Pu.D., Sc.D. (Read April 19, 1918.) The present enumeration of the mussels of the upper Tennessee is the result of the writer’s work carried on in this region since 1912, under the auspices of the Carnegie Museum of Pittsburgh, Dr. W. J. Holland, director. It is intended, herein, to give a complete synopsis of this fauna, using the modern system, and the accepted rules of nomenclature, together with a full synonymy of the various forms, as far as it has been firmly established. Much stress has been laid upon the facts of the geographical distribution, because it has become evident that not all of the species are uniformly distributed in this area. The latter includes all of the upper Tennessee drainage from Chattanooga, Tenn., upward, comprising largely eastern Tennessee, a small section of northern Georgia, and parts of North Carolina (in the high mountains), and southwestern Virginia. It appears that the Walden Gorge of the Tennessee River, below Chattanooga, forms some kind of a barrier to Nayad distribution, at least for cer- tain species; at any rate, it forms a natural division within the Ten- nessee system. Of course, not all parts of this drainage have been investigated by myself; but collections have been made in all of the more important streams; and, together with the records obtained from other sources, it is believed that this fauna is now rather com- pletely known. The region in question is known as one of the chief centers of Nayad development, and may be called the most prolific section of the world in this particular group. Many of the species described by the older authors (Conrad, Lea, and others) originally came from this region. But no synopsis of the whole fauna has been pub- lished, except the attempt made by Lewis (1871 and 1872). Reprinted from Proceedings American Philosophical Society, Vol. lvit., 1978 522 ORTMANN—NAYADES OF In this connection it should be noted that Lewis’s ‘‘ Holston ’ River” actually is the Tennessee River below Knoxville (chiefly in the region of Concord). Indeed, even at present, the natives in Con- cord call the river there “ Holston”; but the maps of the U. S. Geo- logical Survey give the name Tennessee to the river below the junc- tion of the Holston (proper) and French Broad. In Knoxville the river is called Tennessee. Aside from Lewis’s paper, only a few are at hand which con- tribute to the fauna of this region. One of them has been published by Pilsbry and Rhoads (1896). This is, however, by no means a synopsis, containing hardly half of the forms which are found here. But on account of the good locality-records it is very valuable; in fact, it is, up to the present time, the most accurate publication in this respect. And further, two preliminary papers have been pub- lished, based upon my own collections; one by myself (Ortmann, 19130), the other by Goodrich (1913). . These, however, treat only of the headwaters-region of Powell, Clinch, and Holston in Virginia. In addition to the material collected by myself, I have examined the upper Tennessee shells in the collection of Mr. B. Walker in De- troit, and I want to express to Mr. Walker my best thanks for the privilege of examining his shells, and the delightful days I spent in his home in April, 1916. Mr. Walker has a great number of shells from this region obtained from older collections, which in part are cotypes, topotypes, or other authentic material. But the greatest treasure in his collection are the Nayades collected by Professor Dr. C. C. Adams in 1899 to 1901, in the course of his work on Jo, be- cause Professor Adams always was very careful in recording his localities. ‘ A large number of the “species” described by Lea (generally from very insufficient material), and of those listed by Lewis, are synonyms. Additional species have been subsequently described by various authors; but also these are mostly synonyms. There is a rule, observed in many cases, and indicated first by Wilson & Clark, 14, that one and the same shell assumes different shapes in the large rivers and in small streams and headwaters, a rule the existence of which will be shown‘elsewhere; and it is easily understood why the various local races have been regarded as good species, as long as 523 UPPER TENNESSEE DRAINAGE, a me a =: 4 do) | { f as ob oO ° oL ° Ae e = £ e 3 ; ; . ean z p i é Ne coke %, Z iy \ -——_—— ono ae == anne adeeb N= Pie ee . ; een ; ; ae : . ve Kda¥nk 4 a Vo e=4 ee --"7 = ~——? ITNABIINIg > wa) ~ 440390149 e- GZS go! . we 3) alk > iw a) vse “af 3 ot AS, Os lek NYY Aue y EOLA QZITIAN3Y ABN SSAwoHe 7 wre se $e AW A280 LG ; & ” As ty 2 x «lf 033N, i aie e . oe , RS VA 1V One, c wey aS 3 iz a8 4 So 7 by, %, a Pran0d GO rats of a Pt ? %, ‘ 556 ORTMANN—NAYADES OF Everywhere in the larger rivers: Tennessee, Holston, French Broad, Clinch, and Powell; goes up, in the Powell, to Jonesville, Lee Co., Va.; in the Clinch, to Clinchport, Scott ‘Co., Va.; in the Holston to the South Fork at Pactolus, Sullivan Co., Tenn.; in the French Broad, it goes to the Nolichucky at its mouth( Chunn’s Shoals, Hamblen Co., Tenn.). It is also in Emory River, up to Harriman Junction, Roan Co., Tenn., and in Hiwassee River, at Kincannon Ferry, Meigs Co., Tenn. In the Tennessee below Knox- ville, and down to Chattanooga, it is extremely abundant. Type locality: Ohio River. 34. ELLIPTIO DILATATUS (Rafinesque), 1820. Unio dilatatus Rafinesque, ’20——Unio gibbosus Lewis, ’71—Unio gibbosus Pilsbry & Rhoads, ’96.—Elliptio gibbosus Ortmann, '12b, p. 271 (anatomy ).—Elliptio dilatatus Ortmann, 7130, p. 311. —Elliptio gibbosus Goodrich, ’13, p. 94.—Unio gibbosus Simpson, 14, p. 597.—Unio dilatatus Vanatta, ’15, p. 555.—Ellptio dilatata Utterback, ’16, p. go. Common, in large rivers as well as in small creeks, possibly the most widely distributed species in the upper Tennessee region, so that it is heardly required to name special localities. However, it should be mentioned that it is one of the species which go up, in French Broad River, to Asheville, Buncombe Co., N. Car. (Walker coll.). , In our region, this species is of moderate size, and purple color of the nacre prevails; only specimens from the French Broad are more frequently white-nacred. Type locality: Ohio River (the type is from Kentucky River, ac- cording to Vanatta). Genus: LASTENA Rafinesque (1820). Ortmann, ’12), p. 297, and ’15, p. 106. 35. LASTENA LATA (Rafinesque), 1820. Anodonta lata Rafinesque, ’20.—Margaritana dehiscens Lewis, ’71.— Lastena lata Goodrich, ’13, p. 94—Lastena lata Ortmann, ’I5, p. 106 (anatomy ).—Lastena lata Vanatta, ’15, p. 554. UPPER TENNESSEE DRAINAGE. 557 Reported from Tennessee River, below Knoxville (Lewis), and the Holston River (Call). I have never found it in the Holston, but only in the Clinch, at Edgemoor and Clinton, Anderson Co., Tenn. ; at Oakman, Grainger Co., Tenn. In the Walker collection it is from Clinch River at Clinchport, Scott. Co., Va.; and I collected it still farther up at St. Paul, Wise Co., and Cleveland, Russel Co., Va. It is undoubtedly a rare shell. Type locality: Kentucky River. Subfamily: ANODONTINZE Ortmann. Ortmann, IQIO, p. 117. Genus: LAsmMiconaA Rafinesque (1831). Symphynota Lea (1829), Ortmann, ’12b, p. 280.—Lasmigona Frier- Son, 140; p. 40. 36. LASMIGONIA (SULCULARIA) BADIA (Rafinesque), 1831. Alasmodon badium Rafinesque, ’31—Margaritana holstonia Lewis, *71—Symphynota holston(ia) Goodrich, ’13, p.94.—Alasmidonta holstonia Simpson, *14, p. 502—Symphynota (Sulcularia) badia Frierson, ’14a, p. 7——Symphynota (Alasminota) holstonia Ort- mann, ’14, p. 43 (anatomy). The subgenus Alasminota Ortmann, *14, p. 42, is synonym to Sulcularia Rafinesque, ’31 ; see Frierson, |. c. A characteristic small stream species, abundant locally, and re- ported by Lewis from small streams in Monroe Co., Tenn., and by Call from tributaries of the Holston in East Tennessee. The largest streams where I have seen it are the upper Holston proper at Church Hill, Hawkins Co., Tenn., and the Hiwassee, at Austral, Polk Co., Tenn. In both cases the specimens came from a small slough. In the headwater streams, it is in upper Powell, in Va., in the uppermost Clinch in Tazewell Co., Va.; in Little Mocassin Creek, Scott Co., Va.; South Fork Holston at Bluff City, Sullivan Co., Tenn.; Watauga River, Carter Co., Tenn. Other small streams are the following: Cove Creek, Campbell Co., Tenn. (to Clinch) ; Bull Run, Knox Co., Tenn. (to Clinch) ; 558 ORTMANN—NAYADES OF Big Creek, Hawkins Co., Tenn. (to Holston) ; Long Creek, Cocke Co;, Lenn. (to French Broad); Little Piceon’ River, Sevier (Gos Tenn. (to French Broad) ; First Creek and Third Creek, Knox Co., Tenn. (to Tennessee); Piney River, Rhea Co., Tenn. (to Ten- nessee) ; Conasauga .Creek, Monroe Co., Tenn. (to Hiwassee) ; South Chickamauga Creek, Catoosa Co., Ga. (to Tennessee). Thus this species has practically a universal distribution in our region, but it strictly avoids larger streams. Type locality: Small streams of the Knobs, Kentucky (head- waters region of Cumberland and Kentucky Rivers). 37. LASMIGONA (LASMIGONA) COSTATA (Rafinesque), 1820. Alasnudonta costata Rafinesque, ’20.—Margaritana rugosa Lewis, *71.—Alasmodonta rugosa Pilsbry & Rhoads, ’98—Symphynota costata Ortmann, ’12b, p. 283 (anatomy ).—Symphynota costata Ortmann, ’130, p. 311.—Symphynota costata Goodrich, ’13, p. 94. —Symphynota (Lasmigona) costata Simpson, ’14, p. 488. A common species, most abundant in small and medium-sized streams, rarer in the larger rivers, but not absent there (Tennessee, lower Clinch and Holston). Goes up, in the Powell, to Olinger, Lee Co., Va.; in the Clinch, to Cedar Bluff, Tazewell Co., Va.; it is in Emory River; goes, in North Fork Holston, to Saltville, Smyth Co., Va.; in Middle Fork Holston, to Chilhowie, Smyth Co., Va.; in South Fork Holston, to Barron, Washington Co., Va.; in Watauga, to Johnson City, Washington Co., Tenn. It is also in South Chicka- mauga Creek, Ringgold, Catoosa Co., Ga. Type locality: Kentucky River. Genus: ANoponTA Lamarck (1799). Ortmann, ’12), p. 286. 38. ANODONTA GRANDIS GIGANTEA (Lea), 1834. Anodonta gigantea Lea, ’34.—Anodonta grandis (incl. gigantea) Ortmann, 12), p. 292 (anatomy).—Anodonta grandis gigantea Simpson, ’14, p. 420. No Anodonta has ever been reported from the upper Tennessee UPPER TENNESSEE DRAINAGE. 559 region. However, in the collection of B. Walker, there are two large specimens of a form of Anodonta grandis Say, collected by Mr. M. D. Barber, of Knoxville, in a “small mud pond near French Broad River, 8 miles above Knoxville.” The largest has a length of 168 mm., the other of 127 mm. In the latter, the beak sculpture is vis- ible, and corresponds to that of A. grandis; it has white nacre, and much resembles specimens of the var. gigantea Lea, as found in western Pennsylvania. The other (larger) is a little distorted and tapering behind, and has pale purple nacre, resembling in these char- acters some of the southern forms of the species. The occurrence of this form in this isolated locality is quite re- markable, but supports the view that Anodontas may possess excep- tional means of dispersal. According to more detailed information obtained by Mr. Walker in 1916 from Mr. Barber, the pond is “plainly natural, some 4 or 5 rods long by 2 rods wide, with soft deep mud. There was a small stream of water running to the French Broad River, just a short distance. This was 10 or II years ago.’ “Two years ago, I took a nephew and found the same pond, but although we waded the pond in every direction, up to our knees in mud, we could not find even a fragment of gigantea. A man liv- ing near there said he had seen large shells in another pond near, but I have not examined it.” These ponds, a short distance from the river, probably are on the flood plain of French Broad; the writer has not been able to locate them on the U. S. Geol. Surv. maps. Type locality: Port Gibson, ‘Claiborne Co., Miss. Genus: ANODONTOIDES Simpson (1898). Ortmann, 120, p. 294. 39. ANODONTOIDES FERUSSACIANUS (Lea), 1834. Anodonta ferussaciana Lea, ’34.—Anodontoides ferussacianus Ort- mann, *12b, p. 294 (anatomy)—Anodontoides ferussacianus Simpson, ’14, p. 467. Anodonta oblita (?) Lewis, ’71, may possibly stand for this species. Lewis doubtfully reports this from the Tennessee below Knoxville, but this probably is a mistake. Originally, 4. oblita Lea 560 ORTMANN—NAYADES OF and Anodonta demgrata Lea, which are this species, have been de- scribed from Campbell Co., Tenn.: this is surely in the Cumberland drainage, since Wilson & Clark report it (14) from Clear Fork, at Jellico, Campbell Co., Tenn., and other places in the upper Cumber- land region in Kentucky (I found it myself in Cumberland River at Orby, Bell \Co., Ky.)... In the Walker collection are two specimens from Powell River, Lee Co., Va., collected by G. F. Sword. There is no mistake about them, and thus this species must be listed with the upper Tennessee shells, although known only from a single and somewhat indefinite, locality, and remarkable for its absence all over the rest of this region. Type locality: Ohio River, Cincinnati, Ohio. Genus: ALASMIDONTA Say (1818). Ortmann, ’12), p. 294. 40. ALASMIDONTA (PRESSODONTA) MINOR (Lea), 1845. Margaritana minor Lea, ’45.—Margaritana minor Lewis, °72.— Alasmidonta (Pressodonta) minor Ortmann, ’12b, p. 295.—'14, p. 46 (anatomy ).—Alasmidonta minor Ortmann, 713), p. 311.— Alasmidonta minor Goodrich, ’13, p. 94.—Alasmidonta (Presso- donta) minor Simpson, 14, p. 408. A characteristic small creek species, locally abundant. It is found all over the region, but strictly avoids the medium-sized and larger rivers. JI have never seen it in the Clinch South of the Va.-Tenn. state line, and never in the Holston proper. I know it from the fol- lowing stations. South Fork Powell River, Big Stone Gap, Wise Co., Va.; North Fork Clinch River, Tazewell, Tazewell Co., Va. (Walker coll.) ; Clinch River, Cedar Bluff, and Richland, Tazewell Co., Va.; Clinch River, Cleveland, Russell Co., Va.; Clinch River, St. Paul, Wise Co., Va.; Clinch River, Speers Ferry, Scott Co., Va.; Brush Fork (tribu- tary to Poplar Creek and Clinch), Marlow, Anderson Co., Tenn. ; Big Mocassin Creek, Mocassin Gap, Scott Co., Va.; North Fork Hol- ston River, Saltville, Smyth Co., Va.; Middle Fork Holston River, UPPER TENNESSEE DRAINAGE. 561 Chilhowie, Smyth Co., Va.; South Fork Holston River, Barron, Washington Co., Va.; Big Creek (tributary to Holston), Rogerville, Hawkins Co., Tenn. (Walker coll.) ; Little Pigeon River, Sevier- ville, Sevier Co., Tenn.; Boyd Creek, Boyd Creek, Sevier Co., Tenn. ; Little River, Melrose, Blount Co., Tenn.; Pistol Creek, Rockford, Blount Co., Tenn.; Conasauga Creek, Monroe Co., Tenn. (Walker coll.) ; South Chickamauga Creek, Ringgold, Catoosa Co., Ga. Type locality: Tennessee (and “ North Carolina,’ but no exact locality given). 41. ALASMIDONTA (DECURAMBIS) MARGINATA (Say), 1819. Alasmodonta marginata Say, ’19.—Margaritana marginata Lewis, *71—Alasmodonta marginata Pilsbry & Rhoads, ’96.—Alasmi- donta marginata Ortmann, ’12), p. 297 (anatomy ).—Alasmidonta marginata Ortmann, 7130, p. 311—Alasmidonta marginata Good- rich, 713, p. 94.—Alasmidonta (Rugtfera) marginata Simpson, 14a Ds SOA: As to the subgenus Decurambis Raf., ’31, see Frierson, ’14a, p. 7. Generally distributed over the whole upper Tennessee region, but apparently more abundant toward the headwaters. Goes, in the Powell, to Olinger, Lee Co., Va.; in the Clinch, to Richland, Taze- well Co., Va.; in the Forks of the Holston, to Saltville and Chilhowie, Smyth Co., Va., and is also in Big Mocassin Creek, Scott Co., Va. Pilsbry & Rhoads report it from Watauga River at Johnson City, Washington Co., Tenn. It is also in Ocoee River, at Ducktown, Polk Co., Tenn. (Walker coll.). Type locality: Scioto River, Chillicothe, Ohio. 42. ALASMIDONTA (DECURAMBIS) RAVENELIANA (Lea), 1834. Margaritana raveneliana Lea, ’34.—Alasmidonta (Rugifera) ravene- liana Simpson, 14, p. 507. Differs from A. marginata chiefly in the absence of rugosities on the posterior slope. However, in very young specimens, traces of them are sometimes seen, so that this characteristic feature of the subgenus Decurambis (Rugifera) is still indicated. This form un- doubtedly is an offshoot of the marginata-stock, separated from the 562 ORTMANN—NAYADES OF rest in the high mountains of North Carolina, and developed there into a good species. The type locality is French Broad and Swananoa rivers, Ashe- ville, Buncombe Co., N. Car. I have not been able to find this species there, for the French Broad is polluted in this region (lumber indus- tries on Davidson River). But I have rediscovered it in Big Pigeon River, at Canton, Haywood Co., N. Car., where it is not rare at the proper places. Specimens from “North and South Fork of the Cumberland River,” referred to this species, do not belong here, but are a form of A. marginata, with well-developed rugosities upon the posterior slope (atropurpurea Raf., ’31). Genus: PectAs Simpson (1900). Ortmann, ’14, pp. 45 and 65 (as subgenus). On account of the very peculiar glochidia I consider it better to retain Pegias as a genus, closely allied to Alasmidonta. 43. PEGIAS FABULA (Lea), 1836. Margarita (Margaritana) fabula Lea, ’36—Alasmidonta fabula Ortmann, ’130, p. 311—Alasmidonta (Pegias) fabula Ortmann, "14, p. 65 (anatomy).—Pegias fabula Simpson, ’14, p. 473. A rare species in the upper Tennessee drainage, apparently pre- ferring smaller streams. Possibly it has been often overlooked on account of its small size and the peculiarity of being generally much eroded. I know it from the following localities. Wallen Creek, Lee Co., Va. (Walker coll.) ; Powell River, Dry- den, Lee Co., Va.; Big Mocassin Creek, Mocassin Gap, Scott Co., Va.; North Fork Holston River, Saltville, Smyth Co., Va.; North Fork Holston River, Holston, Washington Co. (Walker coll.) and Mendota, Washington Co., Va.; South Fork Holston River, Pac- tolus, Sullivan Co., Tenn. Type locality: Cumberland River, Tenn. Genus: StropHiITus Rafinesque (1820). Ortmann, ’120, p. 299. UPPER TENNESSEE DRAINAGE. 563 44. STROPHITUS EDENTULUS (Say), 1829. Alasmodonta edentula Say, ’29.—Anodonta edentula Lewis, ’72.— Alasmodonta edentula Pilsbry & Rhoads ,’96.—Strophitus eden- tulus Ortmann, ’12b, p. 299 (anatomy).—Strophitus edentulus Ortmann, 713), p. 311.—Strophitus edentulus Goodrich, ’13, p. 94.—Strophitus edentulus Simpson, ’14, p. 345. Walker thinks that the upper Tennessee-form might be distin- guished from the normal edentulus as var. shaefferianus (Lea). He believes that the latter is more compressed and more projecting an- teriorly, and has more frequently reddish nacre. In the average, this appears to be correct, yet there are many specimens in my rich mate- rial which do not exhibit these characters, and cannot be distin- guished from specimens of the normal edentulus, as found, for in- stance, in western Pennsylvania. Thus I do not think it advisable to separate the two forms. This species is abundant, both in larger rivers and smaller streams: in the Tennessee, in the lower French Board, the Holston and its forks and tributaries, and all over the Clinch and Powell drainages. It goes up, in the Powell, to Big Stone Gap, Wise Co., Va.; in the Clinch to Cedar Bluff, Tazewell Co., Va.; in the Forks of the Holston, to Saltville, Smyth Co., Va., and Bluff City, Sullivan Co., Tenn. It has not been found by myself in the eastern tributaries of the Tennessee south of the French Broad, but possibly this is accidental. Type locality: Wabash River. Subfamily: LAMPSILINZ® Ortmann, 1910. Ortmann, ’10, p. 118. Genus: ELLipsarIA Rafinesque (1820). Ptychobranchus Simpson, ’0o.—Ortmann, 12), p. 305.—Ellipsaria Prierson. 14d, p. 7. 45. ELLIPSARIA FASCIOLARIS (Rafinesque), 1820. Obliquaria fasciolaris Rafinesque, ’20.—Unio phaseolus Lewis, ’71I. —Unio phaseolus Pisbry & Rhoads, ’96.—Ptychobranchus 564 ORTMANN—NAYADES OF phaseolus Ortmann, ’12), p. 306 (anatomy ).—Ptychobranchus phaseolus Goodrich, ’13, p. 94—Ptychobranchus phaseolus Simp- son, “14, p. 333.—Ellipsaria fasciolaris Frierson, ’I4a, p. 7.— Ptychobranchus fasciolaris Vanatta, ’15, p. 554. Widely and uniformly distributed over the upper Tennessee region, but nowhere in great numbers. In the Tennessee, French Broad, Holston, Clinch, lower Emory, and Powell. It goes up, in the Powell, to Pennington Gap, Lee Co., Va.; in the Clinch, to Cleve- land, Russell Co., Va.; in North Fork Holston, to Mendota, Wash- ington Co., Va.; in South Fork Holston, to Emmett, Sullivan Co., Tenn. It is one of the species which has been reported from French Broad River, at Asheville, Buncombe Co., N. Car. It has not yet been found in Little River, Little Tennessee, and Hiwassee, but it is in South Chickamauga Creek, at Ringgold, Catoosa Co., Ga. Type locality: “ Ohio, Wabash, Kentucky rivers.” (The type is from Kentucky River, according to Vanatta.) 46. ELLIPSARIA SUBTENTA (Say), 1825. Unio subtentus Say, ’25——Unio subtentus Lewis, ’71—Unio sub- tentus Pilsbry & Rhoads, ’96.—Ptychobranchus subtentus Ort- mann, *12b, p. 308 (anatomy ).—Ptychobranchus subtentus Ort- mann, “130, p. 311—Ptychobranchus subtentus Goodrich, ’13, p. 94.—Ptychobranchus subtentum Simpson, TAs Passo: Known from Tennessee, Powell, Clinch, Holston, and Nolichucky rivers, but more abundant toward the headwaters, and rather rare in the big rivers.. Goes up to Big Stone Gap, Wise Co., Va.; to Cedar Bluff, Tazewell Co., Va.; and to Smyth Co., Va. (in North and Middle Fork Holston). Also in Big Mocassin Creek, in Scott Co;, Va." Thusat ascends, in the smal] streams, father than 2- fasciolaris. In the headwaters it is locally quite abundant. Type locality: North Fork Holston River (Say says in South Carolina, but this is in Virginia) (topotypes examined). Genus: OBLIQUARIA Rafinesque (1820). Ortmann, ’120, p. 300. UPPER TENNESSEE DRAINAGE. 565 47. OBLIQUARIA REFLEXA Rafinesque (1820). Obliquaria reflera Rafinesque, ’20.—Unio cornutus Lewis, ’71.— Unio cornutus Pilsbry & Rhoads, ’96.—Obliquaria refleva Ort- mann, 7120, p. 310 (anatomy) —Obliquaria reflexa Simpson, ’14, P. 330. Only in the larger rivers. Tennessee below Knoxville (Lewis), and at Rathburn, Hamilton Co., Tenn. (Walker coll.). Lower Clinch, Patton’s Ferry, Roan Co., Tenn. (Pilsbry & Rhoads). I found it in the Clinch at Solway, Knox Co.; Edgemoor and Clinton, Anderson Co., Tenn. In the Walker coll. is a specimen from below Agee, Campbell Co., Tenn. Type locality: Kentucky River and Letart Falls (according to Vanatta, the types are from the latter place, below Parkersburg, Wide) Genus: CypROGENIA Agassiz (1852). Ortmann, 120,-p. 312. 48. CYPROGENIA STEGARIA (Rafinesque), 1820. Obovaria stegaria Rafinesque, ’20.—Unio irroratus Lewis, ’71.— Unio trroratus Pilsbry & Rhoads, ’96.—Cyprogenia irrorata Ort- mann, ’12), p. 312 (anatomy).—Cyprogenia irrorata Simpson, "14, p. 320—Cyprogema stegaria Vanatta, 15, p. 554. Known from the Tennessee below Knoxville (Lewis), and the Holston, at Boyd Island, Knox Co. (Pilsbry and Rhoads). It is also in the Tennessee at Rathburn, Hamilton Co., Tenn. (Walker collec- tion). I traced it up, in the Holston, to Turley Mill, Grainger Co., Tenn. ; and in the lower Clinch it is quite abundant, going up to Oak- man, Grainger Co., Tenn. Type locality: Ohio River. Genus: Dromus Simpson (1900). Ortmann, ’120, p. 314. 566 ORTMANN—NAYADES OF 49. DromMus proMAs (Lea), 1834. Unio: dromas Lea, ’34.—Unio dromas Lewis, ’71—Umnio dromas Pilsbry & Rhoads, ’96.—Dromus dromas Ortmann, ’12), p. 315 (anatomy ).—Dromus dromas Simpson, ’14, p. 341. The typical form is rather swollen, and has a large knob or hump ‘on each valve. This knob, however, is very variable. This form is found in the Tennessee proper. It has been re- ported by Lewis from below Knoxville, and by Pilsbry and Rhoads from Chattanooga, Hamilton Co. It is in the Walker collection from Chattanooga and from Rathburn, Hamilton Co. Pilsbry and Rhoads report it also from the Holston, at Boyd Island, near Knoxville. I found it only in the Tennessee, three miles below Knoxville. There are occasional specimens, which might be called by this name, in the lower Clinch and Holston, but in this region it is gen- erally replaced by the next form, with which it intergrades. Type locality: Harpeth River, Tenn. (and Cumberland River, Nashville, Tenn.). 50. DROMUS DROMAS CAPERATUS (Lea), 1845. Unio caperatus Lea, ’45—Unio caperatus' Lewis, ’71—Dromus caperatus Simpson, ’14, p. 343. This form has lost the “hump,” and it is more compressed than the normal form. It begins in the Tennessee at Knoxville (Lewis), and ascends both the Clinch and Holston, where it intergrades, in the lower parts, with typical D. dromas. In the Holston, it goes up to Holston Station, Grainger Co., Tenn., and in the Clinch, it goes to Clinch River Station, Claiborne Co., Tenn. It enters also the Powell, and goes up to Shawanee, Claiborne Co., Tenn. It is quite abundant in the Holston in Knox and Jefferson Cos., although it is frequently associated here with the typical form, and intergrades with it. Its metropolis, however, is in the Clinch from Anderson Co. upward, and in the Powell, where it is a pure race. Type locality: Clinch River (topotypes examined). Note: The tendency to develop here a more compressed form in the smaller rivers agrees entirely with the similar phenomenon ob- served in species of Fusconaia, ete. UPPER TENNESSEE DRAINAGE. 567 Genus: OxsovariA Rafinesque (1820). Ortmann, 12), p. 320. 51. OBOVARIA (OBOVARIA) RETUSA (Lamarck), 1819. Unio retusa Lamarck, ’19—Obovaria retusa Ortmann, 12), p. 321 (anatomy ).—Obovaria (Obovaria) retusa Simpson, 14, p. 290. Reported by Call from the Holston River in east Tennessee, but missing in Lewis’s list. There are specimens in the Carnegie Mu- seum (from the Smith collection) labeled: Tennessee River, Knox Co., Tenn.; and others in the Walker collection labeled: Holston River, Knox Co., and Holston River, Knoxville, Tenn. In all these cases, apparently, the Tennessee River at and below Knoxville is meant. I found this species only once: a young specimen in Clinch River, at Clinton, Anderson Co., Tenn. In the Walker collection are also specimens from the Tennessee at Bridgeport, Jackson Co., Ala., and from Florence, Lauderdale Co., Ala. (also reported by Hinkley). Thus this species seems to belong to the upper Tennessee fauna, going up to Knoxville and into the lower Clinch; but it apparently is very rare. Some of my specimens of O. subrotunda from the Holston (Mascot) have slighly incurved beaks and purple nacre, and resemble O. retusa to a dergee. Already Wilson and Clark (’14) have indi- cated a similar approaching of the two species in Cumberland River. Type locality: ? (Nova Scotia per errorem). 52. OBOVARIA (OBOVARIA) SUBROTUNDA (Rafinesque), 1820. Obliquaria subrotunda Rafinesque, ’20—Unio circulus Lewis, ’71.— Unio circulus Pilsbry & Rhoads, ’96—Obovaria circulus Ort- mann, ’12b (anatomy ).—Obovaria (Obovaria) circulus Simpson, "14, p. 291 —Obovaria subrotunda Vanatta, ’15, p. 552. The identity of Obliquaria subrotunda and Obovaria striata Rafinesque with U. circulus Lea has been recognized by Conrad in 1834, who selected swbrotunda as name, which thus must be used. Apparently rare in the upper Tennessee region. Reported from the Tennessee below Knoxville (Lewis), and at Knoxville (Call, 568 ORTMANN—NAYADES OF and Pilsbry & Rhoads). I know it only from the Holston River at Mascot, Knox Co., where I found three specimens with the diameter of 60, 61 and 62 per cent. of the length. These must be placed with typical swbrotunda (diameter 60 per cent. and over). A fourth specimen, found associated with these has the diameter of 55 per cent., and should be called O. subrotunda levigata (which see). In this region, there is evidently no tendency of this form to go into the small streams of the headwaters, although my specimens from the Holston indicate an inclination toward the small stream form levigata. Type locality: Ohio River (type from Kentucky River, accord- ing to Vanatta). 53. OBOVARIA (OBOVARIA) SUBROTUNDA LEVIGATA (Rafinesque), 1820. Unio levigata Rafinesque, ’20.—Obovaria (Obovaria) lens Simpson, "14, p. 293.—Obovaria levigata Vanatta, ’15, p. 552. Already Conrad (1834) has seen that Jevigata Raf. is the same as lens Lea. This is the small stream form of the main species, distinguished by greater compression of the shell (diameter less than 60 per cent. of the length). It is quite abundant in the tributaries of the Ten- nessee below the Walden Gorge (Sequatchie River, Tenn., Flint River and Hurricane Creek, Madison Co., Ala., Elk River and Bear Creek). It intergrades here with the main species, as it does in the upper Ohio region. From above the Walden Gorge, I have it from South Chicka- mauga Creek, Ringgold, Catoosa Co., Ga. From the headwaters region, above Knoxville, I have just two specimens from the Hol- ston, one from Mascot, another from Holston Station, Grainger Co., in which the diameter falls under 60 per cent. (to 55 per cent. in the first, to 57 per cent. in the other). I have never seen a trace of this form in the small streams of this region. Type locality: Kentucky River. UPPER TENNESSEE DRAINAGE. 569 Genus: NEPHRONAIAS Fischer & Crosse (1893).* Ortmann, ’12), p. 324. 54. NEPHRONAIAS LIGAMENTINA GIBBA (Simpson), 1900. Unio ligamentinus Lewis, ’71—Unio ligamentinus Pilsbry & Rhoads, ’°96. —Nephronaias ligamentina (incl. gibba) Ortmann, 7120, (anatomy ).—Nephronatas ligamentina gibba Goodrich, ’13, Pp. 94.—Lampsilis ligamentina gibba Simpson, ’14, p. 82. The main species is not represented in the upper Tennessee region, although occasional specimens turn up, which are hard to dis- tinguish from it. The var. gibba is extremely abundant in all the larger rivers: Powell, Clinch, Holston, French Broad, Tennessee, but in the upstream direction it disappears before it reaches the head- waters region. In the Powell, it has been observed up to Shawanee, Claiborne Co., Tenn.; in the Clinch, up to St. Paul, Wise Co., Va.; in the North Fork Holston, up to Holston Bridge, Scott Co., Va.; in the South Fork, up to Pactolus, Sullivan Co., Tenn. From the French Broad it also enters the lower part of the Nolichucky, at Chunn’s Shoals, Hamblen Co., Tenn. Type locality: “Ohio River and southward.” 55. NEPHRONAIAS PECTOROSA (Conrad), 1834. Unio pectorosus Conrad, May, ’34.—Unio perdix Lea, August, ’34. —Unio biangularis Lea, ’40—Unio biangulatus Lea, ’43.—Unio biangulatus Lewis, ’71—Umio biangularis Pilsbry & Rhoads, ’96. —Nephronmas perdix Ortmann, ’12b, p. 3260 (anatomy).— Nephronatas perdix Ortmann, ’13), p. 311.—Nephronaias perdix Goodrich, ’13, p. 94—Lampsilis biangularis and perdix Simpson, "14, pp. 59 and 88. The date of publication of U. perdix Lea is incorrectly given by Simpson as 1827. There is no question that biangularis and perdix are the same species, and it is quite astonishing that this identity has not been recognized by Simpson. Recently, Frierson has suggested to me that vittatis Rafinesque, 1831, from Greene River, Ky., might 1 Possibly the generic name should be changed to Actinonaias; compare Frierson, Nautilus, 31, 1917, p. 48. PROC. AMER. PHIL. SOC., VOL. LVII, LL, OCT. I, 1918. 570 ORTMANN—NAYADES OF be this species. However, I hesitate to accept this, till it has been shown that this species actually exists in Greene River. N. pectorosa is found in the Tennessee below and at Knoxville (Lewis, Pilsbry & Rhoads), but it seems to be rare there. Farther up, it goes into Clinch, Powell, Holston, French Broad, and lower Nolichucky, and it is also in Little River. It ascends toward the headwaters farther than does N. ligamentina gibba, and goes, in the Powell, to Pennington Gap, Lee Co., Va.; in the Clinch to Cleve- land, Russell Co., Va.; in North Fork Holston, to Saltville; Smyth Co., Va.; in South Fork Holston, to Barron, Washington Co., Va.; and in the Watauga, to Watauga, Carter Co., Tenn. Just in the region, where N. ligamentina gibba begins to disappear (north of the Va.-Tenn. state line), N. pectorosa is most abundant and in its best development. Type locality: Elk River, North Alabama. Genus: AMYGDALONAIAS Fischer & Crosse (1893). Ortmann, 120; p. 327. 56. AMYGDALONAIAS TRUNCATA ‘(Rafinesque), 1820. Truncilla truncata Rafinesque, ’20.—Unio elegans Lewis, ’71.— Amygdalonaias elegans Ortmann, ’12b, p. 328 (anatomy ).— Plagiola (Amygdalonaias) elegans Simpson, 14, p. 307.— Plagiola elegans Vanatta, ’15, p. 553.—Amygdalonaias truncata Utterback, ’16, p. 148. Vanatta does not accept Rafinesque’s specific name on account of Unio truncata Spengler, 1793. However, these do not conflict (Walker, ’16). Not abundant, and missing in the headwaters. Tennessee below Knoxville (Lewis) ; Clinch River, Solway, Knox Co.; Edgemoor and Clinton, Anderson Co.; Black Fox Ford, Union Co.; Clinch River Station, Claiborne Co.; Oakman, Grainger Co., Tenn. Hol- ston River, McBee Ford, near Hodges, Jefferson Co., Tenn. Type locality: Ohio River (type from Falls of the Ohio, accord- ing to Vanatta). UPPER TENNESSEE. DRAINAGE. 571 Genus: PLAGIoLA Rafinesque (1820). Ortmann, 120, p. 329. 57. PLAGIOLA LINEOLATA (Rafinesque), 1820. Obliquaria lineolata Rafinesque, ’20.—Unio securis Lewis, ’71.— Unio securts Pilsbry & Rhoads, ’96.—Plagiola securis Ortmann, *12b, p. 329 (anatomy).—Plagiola (Plagiola) securis Simpson, "14, p. 304.—Plagiola lineolata Vanatta, °15, p. 553.—Plagiola securis Walker, ’16, p. 45. Of the names of Rafinesque (depressa, lineolata, and ellipsaria) given to this species, lineolata has been selected by Conrad in 1834. (Walker, 1. c.) This name has been used also by Say, Agassiz, and Call. Only in the larger rivers. Tennessee at and below Knoxville, also at Rathburn, Hamilton Co., Tenn. (Walker collection). In the Clinch at Patton’s Ferry, Roane Co. (Pilsbry &-Rhoads), and at Offutt, Anderson 'Co., Tenn. Rare. Type locality: Falls of the Ohio. Genus: PARAPTERA Ortmann (IQITI). Ortmann, 7120, p. 330. If the first species (leptodon) should actually belong here, the generic name must be dropped in favor of Leptodea Rafinesqtie, ’20 (see Frierson, ’14a, p. 6). Utterback (716, p. 151) uses Lasmonos Raf. 58. PARAPTERA LEPTODON (Rafinesque), 1820. Unio leptodon Rafinesque, ’20.—Unio tenuissimus Lewis, ’71.— Lampsilis (Proptera) leptodon Simpson, 14, p. 188.—Lampsilis leptodon Vanatta, 15, p. 551.—Lasmosos leptodon Utterback, 210, ps di50: A rare shell. Lewis gives it from the Tennessee below Knoxville. In the Walker collection it is from the Clinch, at Needham Ford, Union Co., Tenn., and I found it in the Clinch at Edgemoor, Ander- son Co., Tenn. I found it also in the Holston at Holston Station and Noeton, Grainger Co., Tenn. 572 ORTMANN—NAYADES OF Type locality: Lower Ohio River (Vanatta says that the type is from Kentucky River). 59. PARAPTERA FRAGILIS (Rafinesque), 1820. Unio fragilis Rafinesque, ’20——Umnio gracilis Lewis, ’71.—Unio gracilis Pilsbry & Rhoads, ’96.—Paraptera gracilis Ortmann, ’120, p. 331 (anatomy).—Lampsilis (Proptera) gracilis Simpson, ’14, p. 181.—Leptodea (?) fragilis Frierson, ’14a, p. 7—Lampsilis fragilis Vanatta, ’15, p. 552—Lasmonos fragilis Utterback, ’16, Pp: aus 2s In the larger rivers, not rare. Tennessee River, below Knox- ville (Lewis) and Holston River, Boyd Island, near Knoxville (Pilsbry & Rhoads). In the lower Nolichucky at Chunn’s Shoals, Hamblen Co.; up the Holston to Holston Station, Grainger Co.; in the Clinch up to Clinch River Station, Claiborne Co.; in the Powell River, up to Combs, Claiborne Co., Tenn. Type locality: Ohio River (the type is, according to Vanatta, from creeks in Kentucky). Genus: PRopreRA Rafinesque (1819). Ortmann, ’12), p. 332. 60. PROPTERA ALATA (Say), 1817. Unio alatus Say, °17—Unio alatus Lewis, ’71—Unio alatus Pilsbry & Rhoads, ’96.—Proptera alata Ortmann, ’12), p. 333 (anatomy). —Lampsilis (Proptera) alata Simpson, ’14, p. 162. Common in the larger rivers: Tennessee in Knox Co.; goes up the Clinch to Clinchport, Scott Co., Va.; up the Powell, to Combs, Claiborne Co., Tenn.; it reaches the North Fork Holston at its mouth, at Rotherwood, Hawkins Co., Tenn. It is also in French Broad River, and in the lower part of the Nolichucky at Chunn’s Shoals, Hamblen Co., Tenn. Type locality: ? Genus: ToxoLAsMA Rafinesque (1831). Corunculina (error typ.) Simpson, ’98.—Carunculina Ortmann, 12), P.'3375) 14s 1p. Oe: To.xolasma Frierson, ’144, p. 7. UPPER TENNESSEE, DRAINAGE, 573 61. TOXOLASMA LIVIDUM (Rafinesque), 1831. Unio lividus Rafinesque, ’31—Unio moestus Lea, ’41.—Unio cylin- drellus Lea, ’°68.—Unio glans Lewis, ’*71.—Unio glans Pilsbry & Rhoads, ’96.—Lampsilis (Carunculina) cylindrella and moesta Simpson, 714, pp. 155, 156.—To.xolasma livida Frierson, ’14a, Pp. 7- The upper Tennessee form is not the real U. glans of Lea (’43), as already hinted at by Pilsbry & Rhoads. The latter is more swol- len, and has more inflated beaks, and possibly, it is the big river and lowland form of To.xolasma lividum. What Lea has described as U. moestus (from French Broad River, Tenn.) undoubtedly is this: I have specimens from Little Pigeon River (tributary to French Broad), which are fully identical with moestus. U. cylindrellus Lea (Duck River, Tenn.) is in shape absolutely identical with T. lividum; however, it differs by paler color of epidermis and nacre. Such pale specimens occasionally are found in the upper ‘Tennessee drainage as individual variations. Yet it might be, that elsewhere (Tennessee drainage in northern Alabama, and:Alabama drainage in Alabama and Georgia) this pale form becomes a distinct race. U. pullus Conrad, ’38, from the original locality, Wateree River, S. Car., may be a different species. But it is rather sure that the specimens from ‘Warm Springs, N. Car.” (= Hot Springs, Madi- son Co., N. Car., on the French Broad) are actually T. hividum. There is great variation in the color of the nacre: it may be en- tirely dark purple, or may have (generally) a whitish margin along the edge of the shell; or it may (very rarely) have a pale, yellowish, color. The epidermis is mostly blackish or blackish brown, but it may become pale brown or greenish brown. This. species seems to have a wide range over the upper Ten- nessee drainage, but, on the other hand, it seems to be rather local. I know it from the following localities: South Fork Powell River, ° Big Stone Gap, Wise Co., Va. (Walker coll.) ; Powell River, Jones- ville, Lee Co., Va. (Walker coll.) ; Shawanee, Claiborne Co., Tenn. (Walker coll.) ; Green’s Ford, Union Co., Tenn. (Walker coll.) ; Clinch River, Speer’s Perry, Scott Co., Va.; Emory River, Harri- 574 ORTMANN—NAYADES OF man, Roane Co., Tenn.; North Fork Holston River, at Saltville, Smyth Co. (O. A. Peterson), and at Holston, Washington Co., Va. (Walker coll.) ; Holston Bridge, Scott Co., Va. (Walker coll.) ; Holston River, Rogersville (—Austin Mill), Hawkins Co., Tenn. (Walker coll.) ; Little Pigeon River, Sevierville, Sevier Co., Tenn. ; Pistol Creek, Rockford, Blount Co., Tenn. In addition, it is in Lewis’s list from the Tennessee below Knoxville. Type locality: Rockcastle River, Ky. Its actual presence in Rock- castle River, at Livingston, Rockcastle Co., Ky., has recently been confirmed by Williamson (as U. glans, see: Ohio Naturalist, 5, 1905, p. 311). Wilson & Clark, ’14, do not give it from this place, but I have seen specimens in the Walker coll. from this locality. Note: This form is also present in the tributaries of the Ten- nessee in North Alabama, as has been indicated long ago by Conrad. It also seems to be represented in the Alabama drainage, but from this region it generally goes under the name of corvunculus Lea, ’68. I have a number of specimens of the latter before me, which I cannot distinguish from the upper Tennessee form. Genus: Lem1ox Rafinesque (1881). Frierson, ’t4a, p. 7—Ortmann, 16, p. 39. 62. LeMiIox R1MoSus (Rafinesque), 1831. Unio rimosus Rafinesque,’31.—Unio celatus Lewis, ’71.—Micromya celata Goodrich, ’13, p. 94.—Micromya celata Simpson, ’14, p. 34.—Lemiox rimosus Frierson, ’14a, p. 7—Lemnuox rimosus Ort- mann, ’16, p. 39 (anatomy). This species has been reported from the Tennessee below Knox- ville (Lewis), and from Powell, Clinch, and Holston rivers (Call). It seems to have a wide distribution, but it is found nowhere in great numbers, and is thus a rdre shell. In the Powell, it goes up to Jonesville, Lee Co., Va.; in the Clinch, to St. Paul, Wise Co., Va.; in the North Fork Holston, to Holston, Washington Co., Va. It is found here and there in the Holston proper and the lower Clinch, but I have never seen it in the eastern tributaries of the Holston- Tennessee system. UPPER TENNESSEE DRAINAGE. 575 Type locality: Cumberland River (not reported from the Cum- berland drainage by Wilson & Clark, ’14). Genus: Mepronipus Simpson (1900). Orunann, 120) p. 234-15, po 143: 63. MEDIONIDUS PLATEOLUS (Rafinesque), 1831. Unio plateolus Rafinesque, ’31—Umnio conradicus Lewis, ’71.—Unio conradianus Pilsbry & Rhoads, ’96.—Medionidus conradicus Ort- mann, ’120, p. 335, and ’15, p. 142 (anatomy )—Medionidus con- radicus Ortmann, ’13), p. 311—Medionidus conradicus Good- rich, *13, p. 94.—Medionidus conradicus Simpson, ’14, p. 247. The identity of U. plateolus and conradicus Lea (’34) has been suggested to me by Frierson, and I think, this is correct. Very abundant in the headwaters and in small streams generally, but quite rare in the larger rivers. It has been found practically all over our region, and often goes up into the smallest streams which contain at all Nayades. It is hardly worth while to record single localities, and it suffices to say that it is in the Powell, Clinch, and Holston, and their tributaries; in the French Broad, it goes up to Asheville, N. Car.; it is in Little River, and in South Chickamauga Creek, Ga. From the Tennessee proper, below Knoxville, it has been reported by Lewis. Type locality: Falls of the Cumberland River. (Its existence at this place, below the falls, has been confirmed by Wilson & Clark, 14.) Genus: Eurynta Rafinesque (1820). Ortmann, ’120, p. 336. 64. Eurynia (MicroMYA) FABALIS (Lea), 1831. Unio fabalis Lea, ’31—Umnio fabalis Lewis, ’71.—Eurynia (Mi- cromya) fabalis Ortmann, 12), p. 339 (anatomy ).—Eurynia fabalis Goodrich, ’13, p. 94——Micromya fabalis Simpson, ’14, P- 33. Rather rare, but possibly in part overlooked. Lewis gives it from Tennessee River, below Knoxville, and the Walker collection 576 ORTMANN—NAYADES OF has it from Tennessee River, Little River Shoals, Knox Co., Tenn. I found it chiefly toward the headwaters, in Powell, Clinch, and Hol- ston. In the Powell, it goes up to Combs, Claiborne Co., Tenn. ; in the Clinch, to Cleveland, Russell Co., Va.; in the North Fork Hol- ston, to Hilton, Scott Co., Va.; in the South Fork, to Pactolus, Sul- livan Co., Tenn. It seems to be absent from the eastern tributaries of the system. Type locality: Ohio River. 65. EuryniA (MicroMyA) TRABALIS (Conrad), 1834. Unio trabalis Conrad, ’34.—Eurynia (Micromya) trabalis Ortmann, Extremely rare. I found a single specimen in Clinch River, at Speer’s Ferry, Scott Co., Va.; another one in South Chickamauga Creek, Ringgold, Catoosa Co., Ga.; and three specimens in Hiwassee River, at Austral, Polk Co., Tenn. At the first locality, it was asso- ciated with the next species. The only difference of E. trabalis and perpurpurea is, that the former has white, the latter purple nacre. The specimens from the Hiwassee are exceptionally large, and do not taper so much behind as usual. This species seems to have its metropolis in the Cumberland system (Wilson & Clark, ’14). Call cites it. from Clinch and Powell Rivers in Tennessee, but Lewis does not mention it from the “ Hol- ston”? (= Tennessee), and I have never seen a trace of it in the whole Holston drainage. According to Hinkley, it is rare in the mussel shoals of the Tennessee in Alabama; the Carnegie Museum has one specimen from Paint Rock River, at Paint Rock, Jackson Co., Ala. It may be that my localities in Chickamauga Creek and Hiwassee River are connected with the range in northern Alabama. Type locality: ? 66. Eurynta (MIcroMYA) PERPURPUREA (Lea), 1861. Unio perpurpurea Lea, ’61—Euryma perpurpurea Ortmann, 713), p. 311.—Eurynia purpurpurea Goodrich, ’13, p. 94.—Lampsilis perpurpurea Simpson, *14, p. 133.—Eurynia (Micromya) per- purpurea Ortmann, ’15, p. 63 (anatomy). ’12b, p. 340 (anatomy ).—Lampsilis trabalis Simpson, ’14, p. 132- UPPER TENNESSEE DRAINAGE. 577 This may be only a variety with purple nacre of E. trabalis, but I have not yet seen any intergrades. It is characteristic for the Clinch River, where it is not rare in Virginia (from Speer’s Ferry, Scott Co., up to Cedar Bluff, Taze- well Co.). In addition, it is in the Powell at Olinger, Lee Co., Va. (Walker coll.); in the North Fork Holston, from Rotherwood, Hawkins Co., Tenn., to Mendota, Washington Co., Va.; and in Emory River, Harriman, Roane Co., Tenn. In the Tennessee and Holston proper, and their eastern mountain tributaries, no trace of this species has ever been found. Type locality: Tennessee. 67. EURYNIA NEBULOSA (Conrad), 1834. Unio nebulosus Conrad,’34.—Unio cumberlandianus Lea,’36.—Unio cumberlandicus Lea, ’38—Umio notatus Lea, ’38—Unio glaber Lea, °38—Unio creperus Lea, ’°38—Unio muehlfeldianus Lea, °38.— Unio simus Lea, ’38.—Unio obscurus Lea, ’38.—Unio zeig- lerianus Lea, ’38—Unio amenus Lea, ’40.—Unio fatuus Lea, ’40.—Umnio dactylus Lea, ’40——Umio tener Lea, ’40——Unio regu- laris Lea, ’41.—Unio puniceus Haldeman, ’42—Unio radians Lea, ’°57.—Unio jonesi Lea, ’59.—Unio discrepans Lea, ’60.— Unio scitulus Lea, ’60.—Umnio lingueformis Lea, ’60.—Unio per- pictus Lea, ’60—Unio planicostatus Lea, ’60.—Unio sparus Lea, 68.—Unio dispansus Lea, ’71—Unio glaber, iris, cumberlandi- anus, jonest, sparus Lewis, ’71, and ’72.—Unio muhlfeldtianus Pilsbry & Rhoads, ’96—Eurynia nebulosa, dispansa, and plani- costata Ortmann, ’13), p. 311.—Eurynia nebulosa Goodrich, 13, p. ’94.—Euryma (Micromya) nebulosa Ortmann, 715, p. 64 (anatomy ). Simpson, ’14, has recognized the identity of a great number of these nominal “ species,’ but not of all of them. He gives (pp. 119 and 120) : cumberlandianus (Cumberland R., Tenn.) ; notatus (Cum- berland and Holston R., Tenn.) ; glaber (Holston R., Tenn.) ; radians (Othcalooga Cr., Gordon Co., Ga.) ; jonesi (Euharlee Cr., Ga.) ; dis- crepans (North Alabama) ; scitulus (Tuscumbia, Ala.) ; lingueformis (Columbia, Ga., and French Broad R., Tenn.) ; perpictus (Bull R. 578 ORTMANN—NAYADES OF and Holston R.) ; sparus (Swamp Cr., Whitfield Co., Ga.), and I am fully convinced that all these actually are synonyms of nebulosus. But I believe that there are many others, and on account of the ex- traordinary size of the list of synonyms, I think it advisable to make a few remarks as to these. U. creperus Lea, ’38 (Tennessee), made a synonym of L. iris (Lea) by Simpson (p. 115), is founded upon an old half shell, hardly recognizable, but it may be this. In the Walker collection is a specimen from Clinch R., Va. (Wright), labeled creperus, which is distinctly E. nebulosa, It is a male, has purple nacre, and distinct, rather broad, only slightly interrupted rays. U. dispansus Lea, ’71 (east Tennessee), has been put by Simpson (p. 106) in the synonymy of L. vanuxemensis; 1 think it belongs eres U. puniceus Haldeman, ’42 (Simpson, p. 104) (Holston R., Washington Co., Va.). I have found forms corresponding to the description in the same region (topotypes), and they simply are E. nebulosa with a peculiar reddish-orange nacre. U. obscura Lea, ’38 (Nashville, Tenn.), and U. zeiglerianus Lea, °38 (Cumberland R., Tenn.), made synonyms by Simpson (p. 117) are also this, with rather fine, uninterrupted rays, and purple nacre. U. fatuus Lea, ’40 (Holston R., Tenn.), and U. dactylus Lea, ’40 (Caney Fork R., Tenn.), made synonyms by Simpson (pp. 116, 117), are rather elongated and unusually swollen forms of E. nebulosa, with the rays less developed, and not interrupted. U. planicostatus Lea, ’60 (Tuscumbia, Ala.) (Simpson, p. 117). A strongly compressed male, with rays distinct, rather wide, and little interrupted. Such specimens are frequent, chiefly in the Clinch. U. muehlfeldianus Lea, °38 (Cumberland R., Tenn.) (Simpson, p. 121). According to Simpson, only a single specimen is known, which is undoubtedly this, probably a female. Pilsbry & Rhoads give this also from Watauga R., near Johnson City, Washington Co., Tenn.: near this place (Watauga, Carter Co.), I have found E. nebulosa. U. amenus Lea, ’40 (Holston R., Tenn.) (Simpson, p. 122). A typical female of E. nebulosa. U. tener Lea, ’40 (Big Pigeon R., Tenn.) and U. regularis Lea, UPPER TENNESSEE DRAINAGE. 579 41 (French Broad R., Tenn.), made synonyms by Simpson (pp. 122, 123). There are two specimens (J and 9) in the Walker collection, labeled tener (from French Broad, Asheville, N. Car.). They are rather thin-shelled, have rays, which are fine, and subcontinuous in the male, but somewhat spotted in the female. They undoubtedly are a form of E. nebulosa. U. stmus Lea, ’38 (Cumberland R., Tenn.) (Simpson, p. 123). A male, with strongly developed rays: specimens of this type occur frequently, and are practically identical with U. notatus, admitted by Simpson as a synonym of nebulosus. This tremendous synonym indicates that we have to deal here with a very variable species. The variation concerns first of all the color pattern of the epidermis (rays). In the second line it is shown in the color of the nacre, the shape of the shell, and, of course, some- times the females have been made “species”’ for the reason of their different shape. It is not excluded that additional synonyms may be discovered. E. nebulosa belongs in the affinity of E. iris (Lea), ’30, and has practically the same anatomy. Indeed, some of its forms are hardly distinguishable from £. iris, and it may be that the latter is only the western and northwestern representative of it (that of the Ohio drainage). Thus it is also explained why iris has been recorded for * the upper Tennessee River (Lewis). According to my observations, there is only one species of the ivis-group in the upper Tennessee region. It is generally an elon- gated-elliptical shell, more or less pointed behind in the male, slightly dilated and rounded posteriorly in the female, of a yellowish, green- ish, or brownish color, covered more or less with rays, which nor- mally are well developed, and very often broken up into spots. These rays may be fine or wide (in the female, they are often very broad and distinct on the dilated part of the shell), may cover all of the shell, or only part of it, and may be indistinct or nearly miss- ing. The interior of the shell may be white, or of all shades of salmon, orange, pink, or purple. Although I have tried hard, I have been unable to separate this group of forms into species, and it is also impossible for me to dis- tinguish local races. It is true that sometimes specimens from a cer- 580° ORTMANN—NAYADES OF tain locality, chiefly from some small creeks, show uniform and pe- culiar characters, but this holds good only for short distances. In the longer rivers (Powell, Clinch, Holston) the variation is consid- erable and irregular, apparently without any recognizable rules. The distribution of E. nebulosa extends over the whole of the upper Tennessee region, but the species decidedly favors the head- waters and small streams, and often goes up to the uppermost limit of Nayad-distribution in this region. It is not necessary to give a list of the localities: it is practically everywhere in the Powell, Clinch, Holston, French Broad, Little River, Hiwassee. (It apparently is only accidental that it has not been recorded from the Little Ten- nessee.) It is also in Chickamauga Creek, and it deserves special mention that it goes up, in French Broad, to Asheville, N. Car. In the larger rivers, this species is rare, yet it is present. In addition, E. nebulosa has a wide range not only in the Cumber- land drainage, but also in that of the Tennessee in North Alabama, and it also has invaded the headwaters of the Coosa-Alabama system in northern Georgia and Alabama. It is very singular that Wilson & Clark (714) do not mention it in their paper on the Cumberland shells, although a good number of the synonyms have their type locality in this system. Type locality: Black Warrior River, Ala. 68. EuryniA (MicroMyA) VANUXEMENSIS (Lea), 1838. Unio vanuxemensis Lea, ’°38.—Unio nitens Lea, ’40—Unio um- brosus Lea, ’57 (==umbrans Lea, ’57).—Umio tenebricus Lea, *57.—Unio pybasi Lea, ’58.—Unio fabaceus Lea, ’61—Unio copet Lea, ’68.—Unio pybasi Lewis, ,’71—Unio pybasi and caliginosus Pilsbry & Rhoads, ’96.—Eurynia (Micromya) vanuremensis Ortmann, ’12b, p. 34215, p. 65 (anatomy).—Nephronaias copei and Eurynia vanu.remensis Ortmann, ’130, p. 311—Eurynia vanuxemensis Goodrich, ’13, p. 95—Lampsilis vanuxemensis Simpson, 14, p. 105. Unio dispansus Lea, ’71, placed by Simpson here, belongs to E. nebulosa (see above). Probably there are other synonyms. This shell is shorter and higher than E. nebulosa, has generally a UPPER TENNESSEE, DRAINAGE, 581 dark epidermis with indistinct or no rays, and a deep copper-colored or purplish nacre; rarely the latter is lighter, and even when whitish, it has at least some purple or red. The female is characterized by a very strong dilatation of the postbasal margin, and very often, chiefly in old shells, it has a strong “constriction” behind this dilatation. The distribution of this species is very similar to that of E. nebu- losa, preferring also the small streams, and very often the two species are found associated. Also here it is unnecessary to give a list of localities, and it suffices to state that it is found practically over the whole upper Tennessee region. It should be remarked, however, that I did not find it in the headwaters of the Clinch (above Speer’s Ferry, Va.), but it may have been overlooked here. While locally abundant in smaller streams, it becomes rare in the larger rivers. Also here, local races cannot be distinguished, except that the shell attains, in certain streams, a much greater size than in others. This is the case, for instance, in the Middle Fork Holston in Smyth Co., Va., where exceptionally large specimens (U. cope: Lea) are found, while the North Fork contains a small race. Specimens of larger rivers are also generally rather large and well developed, and often lack, in the 9, the posterior constriction. Such specimens have a resemblance to U. lienosus Conrad, being more drawn out at the posterior end. U. lienosus is a southern form, not found in the Tennessee drainage. There is no doubt that U. caliginosus Lea (—=Nenosus), recorded by Pilsbry & Rhoads from the lower Clinch, is founded upon such specimens. Also this species has a wide distribution in the Cumberland drain- age, the Tennessee drainage in Alabama, and the Coosa-Alabama system, and it is very likely that also the form of the Cumberland, questionably referred to lienosa by Wilson & Clark (’14), is this. Type locality: Cumberland River, Tenn. 69. EuryNiA (EurRyYNIA) RECTA (Lamarck), 1819. Unio rectus Lamarck, ’19—Unio rectus Lewis, ’71—Unio rectus Pilsbry & Rhoads, ’96.—Eurynia (Eurynia) recta Ortmann, ’12), Pp. 344 (anatomy )—Eurynia recta Goodrich, ’13, p. 95—Lamp- silis recta Simpson, ’14, p. 95. Abundant in the larger rivers: Tennessee in Knox Co., lower 582 ORTMANN—NAYADES OF French Broad and lower Nolichucky (Hamblen Co.) ; all the way up the Holston to the North Fork at Rotherwood, Hawkins Co., Tenn. Also in the Clinch up to St. Paul, Wise Co., Va., and the Powell up to Combs, Claiborne Co., Tenn. Type locality: Lake Erie (topotypes examined). Note: The Lake Erie form differs somewhat from that of the central basin. If it should be found to be desirable to express this in nomenclature, FE. recta should be reserved for the lake form, and E. recta latissima (Rafinesque), ’20, should be used for the other. Genus: Lampsixis Rafinesque (1820). Ortmann, ’12), p. 345. 70. LAMPSILIS VIRESCENS (Lea), 1858. Unio virescens Lea, ’58.—Lampsilis virescens Simpson, ’14, p. 93. I am not quite satisfied as to the proper position of this species within the genus Lampsilis. According to external appearance, it seems to be related to L. Juteola (Lam.), although it also has some features, which resemble those of L. teres (Raf.) (=anodontoides Lea). In either case, however, it would be a Lampsilis. The type locality is Tennessee River, Tuscumbia, Colbert Co., Ala., and it has been reported, by Call, from Spring Creek at Tus- cumbia. The Carnegie Museum has it from tributaries of the Ten- nessee in northern Alabama (Paint Rock River, and Bear Creek). It has never been found anywhere else. But I have found a few specimens (all males) in Emory River, at Harriman, Roane Co., Tenn., and in the Walker collection are others from Coal Creek, Anderson Co., Tenn. These two streams are not far apart, and flow from Walden Ridge to the Clinch. Emory River is the only western tributary which completely cuts through Walden Ridge and drains, in its headwaters, a section of the Cumberland Plateau. 71. LAMPSILIS OVATA (Say), 1817. Unio ovatus Say, ’17——Unio ovatus Lewis, ’71—Unio ovatus Pils- bry & Rhoads, ’96—Lampsilis ovata Ortmann, ’12b, p. 350 (anatomy ).—Lampsilis ovata Simpson, ’14, p. 48. UPPER TENNESSEE DRAINAGE. 583 Distinguished by the distinct and sharp posterior ridge, depressed (truncated) posterior slope, and the peculiar, wedge-shaped anterior part of the shell. But these characters gradually pass into those of the variety ventricosa. The typical L. ovata is restricted to the larger rivers, and quite abundant there. It is in the Tennessee, the Little Tennessee, Hol- ston, Clinch, and Powell. In the Powell, it goes up to Shawanee, Claiborne Co., Tenn. ; in the Clinch, to Clinchport, Scott Co., Va. It is the prevailing form in the Holston proper, but does not go into the Forks of the Holston. All along its range, and chiefly above Knox- ville, it is accompanied by the var. ventricosa, and intergrades with it. But at the points just named, it disappears, and leaves the field to ventricosa. Type locality: Ohio River. 72, LAMPSILIS OVATA VENTRICOSA (Barnes), 1823. Unio ventricosa Barnes, ’23.—Lampsilis ventricosa Ortmann, ’12), p. 351 (anatomy ).—Lampsilis ovata ventricosa Ortmann, ’130, Pp. 311.—Lampsilis ovata ventricosa Goodrich, ’13, p. 95 —Lampsilis ventricosa Simpson, ’14, p. 38. According to Vanatta (715, p. 551), the type of Lampsilis car- dium Rafinesque, ’20, is this, and also Conrad (’34) says so. How- ever, this conflicts with Rafinesque’s description, from which it is evident that L. cardium is the female of L. ovata. We have here a case where the “type” does not agree with the original description, and it should be borne in mind that the co-called “types” of Rafin- esque, in the Philadelphia Academy, are not types in the strict sense, but merely “ authentic specimens ” of somewhat doubtful value. In this variety, the posterior ridge becomes indistinct, the pos- terior slope is not excavated, and the anterior part of the shell is not remarkably compressed. Also, the shell is generally less convex. But there are all stages of transition. The upper Tennessee form of ventricosa very rarely has the distinct rays of the corresponding form of the upper Ohio region. L. ovata ventricosa is found associated with the normal L. ovata in the larger rivers, but is less frequent there. It goes, however, 584 ORTMANN—NAYADES OF beyond the upper limit of L. ovata in the headwaters, where it is found in its best development and as a pure race. In the Powell, it goes to Olinger, Lee Co., Va.; in the Clinch, to Cedar Bluff, Taze- well Co., Va.; in North Fork Holston, to Saltville, Smyth Co., Va. (also in Big Mocassin Creek) ; and in the South Fork, to Emmett, Sullivan Co., Tenn. It is in Nolichucky River, at Chunn’s Shoals, Hamblen Co., Tenn. (inclining here toward ovata) ; in Little Pigeon River, Sevierville, Sevier Co., Tenn. ; in Little River, Melrose, Blount Co; Tenn; dnd in Emory River, Hartiman;, Roane: Go.) Tenn; vil have seen it also in Chickamauga Creek, Ringgold, Catoosa Co., Ga. Type locality: Wisconsin River, and Mississippi River, Prairie du Chien, Wis. 73. LAMPSILIS FASCIOLA Rafinesque (1820). Lampsilis fasciola Rafinesque, ’20—Unio multiradiatus and perra- diatus Lewis, ’71.—Unio multiradiatus Pilsbry & Rhoads, ’96.— Lampsilis multiradiata Ortmann, 12), p. 352 (anatomy )—Lamp- silis multiradiata Ortmann, ’13b, p. 311—Lampsilis multiradiata Goodrich, ’13, p. 95 —Lampsilis multiradiata Simpson, ’14, p. 55. Also in this case, I do not follow Vanatta’s (’15, p. 551), de- termination of the “type” of fasciola, but rely on Rafinesque’s de- scription, which indicates a shell of the cardiwm-ovata type, with unequal, flexuous rays, which fits multiradiata lea, but not luteola Lamarck. Moreover, Conrad, in 1836, held the same opinion, also pointing out the undulated rays as the main character of this species. Practically everywhere in the larger rivers as well as in smaller streams, but apparently more abundant toward the headwaters. In the Tennessee, South Chickamauga, Hiwassee, Little Tennessee, Little River. In the French Broad drainage, going up here to Ashe- ville, N. Car., and, in Big Pigeon River, to Canton, N. Car. In the Holston, up to the South Fork at Barron, Washington Co., Middle Fork at Chilhowie, Smyth Co., and North Fork at Saltville, Smyth Co., Va. Also in Big Mocassin Creek, and Watauga River. In the Clinch, it goes up to Cedar Bluff, Tazewell Co., Va.; in the Powell, to the North Fork at Big Stone Gap, Wise Co., Va. Also in the Emory at Harriman, Roane Co., Tenn. Type locality: Kentucky River. UPEER TENNESSEE DRAINAGE. 585 74. LAMPSILIS ORBICULATA (Hildreth), 1828. Unio orbiculatus Hildreth, ’28.—Lampsilis orbiculata Ortmann, 7120, p. 343 (anatomy ).—Lampsilis orbiculata Simpson, ’14, p. 76. Not reported previously from the upper Tennessee region, but in the Walker collection are two specimens labeled ‘‘ Holston R., Tenn.,” which come from the Lewis collection (and thus probably are from the Tennessee). 1 have found myself two specimens in the Clinch, one at Offutt, Anderson Co., the other at Solway, Knox Co., Tenn. This species is also in the Tennessee in North Alabama, at the mussel shoals near Florence (reported by Hinkley, and represented in Walker coll. and Carnegie Mus.). Type locality: Muskingum River, Marietta, Ohio. Genus: TRUNCILLA Rafinesque (1820). Ortmann, 120, p. 354. The subgenera distinguished by Simpson require revision, also with regard to nomenclature. I disregard them for the present, re- marking only that the soft parts furnish good criteria for their defi- nition. 75. TRUNCILLA TRIQUETRA Rafinesque, 1820. Truncilla triqueter Rafinesque, ’20.—Unio triangularis Lewis, ’71.— Unio triangularis Pilsbry & Rhoads, ’96—Truncilla triquetra Ortmann, '12), p. 355 (anatomy ).—Tvruncilla triquetra Simpson, Ape 5. Rather frequent both in larger and smaller rivers, but nowhere in great numbers. Tennessee below Knoxville (Lewis) and at Knoxville (Pilsbry & Rhoads) ; Little River in Knox Co. (Walker coll.) ; lower Nolichucky; Holston River up to the South Fork at Pactolus, Sullivan Co., Tenn., and the North Fork at Mendota, Washington Co., Va.; in the Clinch up to Clinchport, Scott Co., Va.; in the Powell, up to Shawanee, Claiborne Co., Tenn. Type locality: Falls of the Ohio. PROC, AMER. PHIL. SOC., VOL. LVII, MM, OCT, I, 1918. 586 ORTMANN—NAYADES OF 76. TRUNCILLA ARCZFORMIS (Lea), 1831. Unio arceformis Lea, ’31—Umio arceformis Lewis, ’71.—Unio arceformis Pilsbry & Rhoads, ’96.—Truncilla arceformis Simp- son, *I4, p. 12. In the larger and medium rivers: Tennessee, Knox Co., Tenn. ; French Broad at Boyd Creek, Sevier Co. In the Holston locally abundant: Boyd Island, near Knoxville (Pilsbry & Rhoads) ; Mc- Millan, Knox Co.; Mascot, Knox Co.; Gant Island near Straw- berry Plains, Jefferson Co. (Walker coll.); McBee Ford, near Hodges, Jefferson Co.; Turley Mill, Noeton, and Holston Station, Grainger Co.; Austin Mill, Hawkins Co., Tenn. In the Clinch at Clinch River Station, Claiborne Co.; and Oakham, Grainger Co., Tenn. Type locality: “Tennessee River.” Lea (Tr. Amer. Philos. Soc., 1834, p. 86) corrects this, saying that, according to Troost, this species is not in the Tennessee, but only in the Cumberland River (meaning, of course, the Tennessee of northern Alabama). 77, TRUNCILLA INTERRUPTA (Rafinesque), 1820. Obliquaria interrupta Rafinesque, ’20.—Unio brevidens Lea, ’31 (not ’24, as given by Simpson).—Unio brevidens Lewis, ’71—Trun- cilla brevidens Simpson, ’14, p. 7. The identity of Rafinesque’s species with that of Lea has been recognized by Conrad, accepted by Kuester and Reeve, and con- firmed by Vanatta (15, p. 550). But Vanatta does not use the name of interrupta, for reasons which do not hold good, as shown by Walker (16, p. 45). The only objection to iterrupta is that it is given, originally, from Kentucky and Ohio rivers, while it seems to be absent at least from the Ohio. But we must remember that Lea’s brevidens also was originally given from Ohio, corrected subse- quently, ’34, to Cumberland River. . In the larger and medium rivers: Tennessee (Lewis), Holston, Clinch, and Powell. In the Powell, up to Rose Hill, Lee Co., Va.; in the Clinch, up to Clinchport, Scott Co., Va.; in the Holston up to the North Fork at Hilton, Scott Co., Va. Type locality: “ Kentucky and Ohio Rivers.” (Probably incor- rect; type from Ohio River, according to Vanatta.) UPPER TENNESSEE DRAINAGE. 587 78. TRUNCILLA LENIOR (Lea), 1843. Unio lenior Lea, ’43.—Truncilla lenior Simpson, ’14, p. IT. AW fare species: Ihave found it im the Clinch, Speers Ferry, Scott Co., Va.; in North Fork Holston, Rotherwood, Hawkins Co., Tenn. ; in South Fork Holston, Pactolus, Sullivan Co., Tenn.; and in the uppermost Holston proper, Church Hill, Hawkins Co., Tenn. There are specimens in the Walker coll. (from Mrs. Andrews), labeled: Holston River, Knox Co. (probably Tennessee River), but Lewis has not recorded it from these parts. For the rest, it is miss- ing in this region, but it turns up again in the Tennessee drainage in North Alabama: Simpson gives it from Paint Rock River, Wood- ville, Jackson Co., Ala., and the Carnegie Museum possesses quite a number of specimens from this river at Paint Rock, Holly Tree, and Trenton, Jackson Co., Ala. The male of this species may be easily recognized by the fine denticles on the margin of the posterior end; for the rest, it looks like a pale-colored Eurynia nebulosa. Type locality: Stones River, Tenn. (Cumberland drainage; but missing in the list of Cumberland shells published by Wilson & @lark,214)) 79. TRUNCILLA HAYSIANA (Lea), 1833. Unio haysianus Lea, ’33.—Unio haysianus Lewis, ’71.—Unio haysi- anus Pilsbry & Rhoads, ’96.—Truncilla haysiana Ortmann, 712), P- 357 (anatomy) —Truncilla haysiana Ortmann, ’13b, p. 311.— Truncilla haysiana Goodrich, ’13, p. 95.—Truncilla haysiana Simpson, ’14, p. 16. Widely distributed, but always only in small numbers at a given locality. Tennessee in Knox Co. (Lewis) and at Knoxville (Pilsbry & Rhoads) ; Little Tennessee, Coytee, Loudon Co., Tenn. (Walker coll.) ; in the Powell, up to Pennington Gap, Lee Co., Va.; in the Clinch, up to Raven, Tazewell Co., Va.; in the Holston, it goes in the North Fork to Hilton, Scott Co., Va., and in the South Fork to Pactolus, Sullivan Co., Tenn. Type locality: Cumberland River. 588 ORTMANN—NAYADES OF 80. ‘TRUNCILLA STEWARDSONI (Lea), 1852. Umio stewardsom Lea, ’52.—Unio stewardsoni Lewis, ’71.—Trun- cula stewardsoni Simpson, ’14, p. 21. I think that what Walker (‘10a, pl. 3, f. 4) has figured as the db of T. lewisi is actually an old ¢ of stewardsoni. A rare species. The Carnegie Museum has specimens from the Tennessee at Knoxville, and Lewis reports it from this region; and there are also specimens from Clinch River, Clinton, Anderson Co., in the Carn. Mus. I found it myself in the Holston, at McMillan and Mascot, Knox Co., and at Holston Station, Grainger Co., Tenn. Type locality: “Chattanooga River, Tenn.” There is no such river in Tennessee. Generally, Lea’s “Chattanooga River” is the Chattooga River in northern Georgia (tributary to Coosa) ; but in the present instance this cannot be, since this species is not found in the Coosa drainage. 81. TRUNCILLA LEWist Walker (1910). Unio foliatus Lewis, ’71.—Truncilla lewisi Simpson, ’14, p. 20. As stated above, Walker’s figure of the ¢ (’10a, pl. 3, f. 4) prob- ably belongs to T. stewardsont. _ I have found a single small male specimen (soft parts examined !) in the Holston, at Holston Station, Grainger Co., Tenn.,and another, somewhat larger one, in the Powell, at Combs, Claiborne Co., Tenn. These differ from the male of T. stewardsoni by a wider radial fur- row, with the two ridges confining it, more divergent. My speci- mens are not full grown, and even if they should be young females (as Walker suggested after examination of the one from the Hol- ston), they would present to us the shape of the male, as all young females do in the genus Trumcilla, exactly as the soft parts of young females resemble those of the males. The Carnegie Museum has (from the Hartman coll.) two fe- males, labeled: Tennessee River, Knox Co., Tenn. Lewis gives this form (as U. foliatus) from the Tennessee at Little River Shoals, below Knoxville; and Walker reports it from Clinch and Holston rivers in Knox Co. (also from Cumberland River, Burnside, Pulaski Co., Ky., but not found here by Wilson & Clark, ’14). UPPER TENNESSEE DRAINAGE. 589 Thus this is undoubtedly one of the rarest species of Truncilla, and its distribution should be studied more closely. Type locality: Holston River, Tenn. (topotype examined). 82. TRUNCILLA PROPINQUA (Lea), 1857. Unio propinquus Lea, ’57—Unio propinquus Lewis, ’71.—Unio propinquus Pilsbry & Rhoads, ’96.—Tvruncilla propinqua Simp- Son, “Ids 'p. 27. ‘Reported, by Lewis, from Tennessee River, Knox Co., and from the Clinch by Call. Pilsbry & Rhoads give it from the Tennessee at Knoxville, the Holston at Boyd Island, near Knoxville, and from the Clinch at Patton’s Ferry, Roane Co. The Carnegie Museum pos- sesses it from the Tennessee, Knox Co. (Smith coll.),and I found it myself in the Clinch at Edgemoor and Clinton, Anderson Co., Tenn. Type locality: Florence and Tuscumbia, Ala. (topotypes ex- amined). 83. TRUNCILLA TORULOSA (Rafinesque), 1820. Amblema torulosa and gibbosa Rafinesque, ’20.—Unio perplexrus Lewis, ’71.—Truncilla perplexa Simpson, ’14, p. 24.—Truncilla torulosa Vanatta, ’15, p. 550. The identity of torwlosa and gibbosa with perplexus has been in- dicated by Conrad (’34), and he selected the name of torulosa. Al- though he later (’36) uses gibbosus (and so do Agassiz and Reeve), the first selection has to stand. The typical T. torulosa has a radial row of prominent knobs across the middle of the shell. But these knobs vary greatly, and in the upstream direction, they have the tendency to become reduced, finally disappearing, thus passing into the condition seen in the next form. . Restricted to the larger rivers. It turns up in the Tennessee at and below Knoxville (Lewis), and continues down the river, but is nardly found above Knoxville. The Carnegie Museum has three young specimens from near Knoxville (Hartman coll.), which have distinct knobs. In addition, there is a specimen from Chattanooga, Hamilton Co., Tenn. (Juny coll.), which is typical in all respects, 590 ORTMANN—NAYADES OF except that it has salmon-colored nacre; to my knowledge, this color of the nacre is extremely rare. Farther down the Tennessee, at the mussel shoals in Alabama, this species is abundant (Conrad, Hinkley, and Carn. Mus.). Type locality: Ohio River and Kentucky River (the type is from Kentucky River, according to Vanatta). 84. TRUNCILLA TORULOSA GUBERNACULUM (Reeve), 1865. Unio gubernaculum Reeve, Conch. icon. 16. Unio. ’65, pl. 28, f. 146. Reeve’s figure undoubtedly is this form. Simpson (’14, p. 26) makes this a synonym of the var. rangiana (Lea), and it surely is the parallel form to rangiana of the upper Ohio drainage. It differs, however, distinctly by the dark green color of the posterior expan- sion of the female shell. From the typical torulosa, this variety differs by the poorly de- veloped or wanting knobs, and by the rather more compressed shell. This is the headwaters form of torulosa, and begins to take its place in the region of Knoxville. I have it from the Nolichucky, Chunn’s Shoals, Hamblen Co.; here, as also in the lower Holston, faint knobs may yet be present. Farther up, the shell is entirely smooth. In the Holston, it goes up to the South Fork at Pactolus, Sullivan Co., Tenn., and to the North Fork at Holston Bridge, Scott Co., Va. In the Clinch, it goes to Dungannon, Scott Co., Va. It is also in the Powell, up to Shawanee, Claiborne Co., Tenn. Locally, it may be quite abundant. Type locality: ? 85. ITRUNCILLA TURGIDULA (Lea), 1858. Unio turgidulus Lea,’’58 (male)—Unio deviatus Reeve, 64 (fe- male).—Truncilla deviata Walker, ’10b, pp. 77, 78, 81.—Truncilla deviata Simpson, ’14, p. 31. Unio turgidulus has been regarded as the male of the female U. florentinus, but it belongs to the female deviatus. In Walker’s key (10b), this has been indicated by the grouping, but it has not been expressly mentinoned. T. turgidula stands nearest to T. biemarginata (Lea), a species UPPER TENNESSEE DRAINAGE. 591 known from the Tennessee in North Alabama, but which has not yet been recorded from the upper Tennessee. T. turgidula agrees with the latter in the biangulate posterior ridge, but the biangulation is much less pronounced, and the depression or furrow in front of it is less developed. In the female (deviatus), the biangulation is also present, but indistinct, and the furrow is obliterated, being filled by the expansion of the shell. The female resembles, to a degree, that of T. florentina, but has the shell, as Simpson states, more elongated, and has fuller and higher beaks (the latter charac- ters hold also good for the male). This species has been recorded from Cumberland and Tennessee rivers (not recently found by Wilson & Clark, ’14, in Cumberland), and from Duck River, Tenn. (Call), but only one definite locality is known (Florence, Lauderdale Co., Ala.). Hinkley reports it from Shoals Creek, Lauderdale Co., Ala. The Carnegie Museum has it from Bear Creek, in Franklin Co., Ala. I have found it in the upper Tennessee drainage, where it is not rare in the Holston proper from Knox Co. up to Austin Mill, Hawkins Co., Tenn. I found it. also in Emory River, Harriman, Roane Co., Tenn. In the Walker coll. it is represented from the Holston, Rogersville, which is prac- tically the same locality as Austin Mill. Type locality: Cumberland River and Florence, Ala. 86. TRUNCILLA FLORENTINA (Lea), 1857. Unio florentinus Lea, ’57—Truncilla florentina Simpson, ’14, p. 30 (excl. turgidulus). This species has not a biangulate posterior ridge, but this ridge is rounded, and the radial depression in front of it is hardly de- veloped, indicated only by a flattening of the shell. In the female, the posterior expansion of the shell may become very large, and is generally of the color of the rest of the shell, or lighter, but not uni- formly dark green, as in 7. capseformis. By the latter character, by the more strongly developed and more numerous denticulations of the margin of the expansion, and by the greater convexity of the shell, T. florentina is distinguished from capseformis. The male of T. florentina is shorter, higher, and more swollen than that of T. capseformis. 592 ORTMANN—NAYADES OF Known from the Cumberland and Tennessee Rivers, from the latter, however, reported hitherto only from North Alabama. I have found it in the Holston, from Knox Co., up to Holston Station, Grainger Co., Tenn., but not in great numbers. Type locality: Florence, Ala. (and Cumberland River) (topotypes examined ). 87. TRUNCELLA WALKERI Wilson & Clark (1914). Truncilla walkeri Wilson & Clark, ’14, p. 46, pl. 1, f. 1. This is practically a large, compressed T. florentina. It agrees with it in every respect, except that it attains a larger size, and is as compressed as 7. capseformis. It may be only the headwaters form of T. florentina. From T. capseformis it is distinguished by larger size, and by the absence of any dark green tints (except rays) upon the posterior expansion of the female... Also the denticulations on the margin of the expansion are stronger and more numerous. Wilson & Clark do not mention these denticulations: but his specimens seem to have been more or less mutilated in this region. My specimens, belong- ing to the type set, show only traces of them, while the fine material I collected myself in the headwaters of the Holston shows them well developed. The males of 7. walkeri and capseformis are very similar: that of walkeri is possibly somewhat larger and of a lighter color, yellow- ish brown, with light green rays, while that of capseformis is green- ish olive, with dark green rays. But these differences are rather un- certain. However, I was never put to the task of separating them, since I have never found the two species associated. T. walkeri is very local in the upper Tennessee region. I found it only in the South Fork Holston at Emmett, Sullivan Co., Tenn., and at Barron, Washington Co., Va. (Walker has it from Barron) ; and further, I found it in Middle Fork Holston, at Chilhowie, Smyth Co., Va. At the latter place it was not rare. In addition, the Carnegie Museum has it from Flint River and Hurricane Creek at Gurley and Maysville, Madison Co., Ala. There are also specimens at hand from the type locality. UPPER TENNESSEE DRAINAGE. 5938 Type locality: East Fork Stones River, Walterhill, Rutherford Co., Tenn. (cotypes examined). 88. TRUNCILLA CAPSAFORMIS (Wea), 1834: Unio capseformis Lea, ’°34——Unio capseformis Lewis, ’71.—Trun- cilla florentina and capseformis Ortmann, ’12), p. 359 (anatomy). —Truncilla capseformis Ortmann, ’13), p. 311 —Truncilla cap- seformis Goodrich, ’13, p. 95.—Truncilla capsefornus Simpson, MiAn pe 20; The specimen of “ florentina,’ of which I have described the anatomy, was actually a capseformis. Distinguished from the two preceding species by the combina- tion of the following characters: shell rather compressed, with the beaks not much elevated; expansion of the female uniformly dark green, with only small, few, and remote denticles on the margin. There are differences in the soft parts of T. turgidula, florentina, walkeri, and capseformis, which will be discussed elsewhere. T. capseformis is very abundant in the upper Tennessee drain- age, all over the region. It is in the Tennessee below Knoxville, in Powell, Clinch, Holston, Nolichucky, Little Pigeon, and goes up, in the French Broad, to Asheville, N. Car. (Walker coll.). It is also in Little ‘Tennessee, at Coytee, Loudon Co., Tenn. (Walker coll.). In the Powell, it has been traced up to Shawanee, Claiborne Co., Wenn; imgthe Clinch; to Cedar Bluit, Vazewell: Co., Vaz; ine the North Fork Holston, to Mendota, Washington Co., Va. (also in Big Mocassin Creek) ; in the South Fork Holston, it is at Pactolus, Sullivan Co., Tenn., but not farther up. Type locality: Cumberland River. ‘The above enumeration contains nearly all nominal species ever recorded from the upper Tennessee region. However, there are yet two additional ones, which have not been mentioned: Unto apacus Haldeman (1824)—Holston River, Tenn. Pleurobema abacus Simpson, ’14, p. 810. A spurious species, which never has been positively recognized. Specimens under this name in the Lea collection (U. S. Nat. Mus.) 594 ORTMANN—NAYADES OF are, as Simpson states, much like “Pleurobema appressus,” that is to say, Lexingtonia dolabelloides conradi (Van.). I have examined these in Washington. Also specimens in the Walker collection, la- beled abacus, from Flint River, Gurley, Madison Co., Ala., are this. MARGARITANA QUADRATA Lea (1861)—FEastern Tennessee. Symphynota quadrata Simpson, ’14, p. 487. The type is lost. This species never has been recognized. I have the suspicion, from description and figure, that it is identical with Alasmidonta minor (Lea), 1845. LIST OF LOCALITIES, AND OF THE NAYAD-FORMS FOUND AT THEM. The following list is submitted for two reasons: first, to give an idea of the richness of the material upon which this paper is founded ; second, to facilitate, for subsequent collectors, the search for cer- tain forms. There will be a time, not far distant, when the fauna of many of the localities will have deteriorated or disappeared in consequence of stream pollution, and thus it is important to know all the localities where a given form has been found. ‘The exact location of all the collecting stations is given on the accompanying map, so that also a change of local geographic nomenclature will not inter- ferew mee pace. 5235) The localities have been arranged according to river systems, be- ginning in the northwest (Powell), and proceeding downstream and eastward. The smaller streams not belonging to the headwaters, have been placed together at the end of each system. The Ten- nessee proper stands at the end of the list. Forms found intergrading at one locality are connected by braces. In several instances of well-known and easily recognized species, I have not taken home specimens at certain localities, but only seen them (mostly dead shells). These are marked “seen.” The record of this fact always was made in the field with actual specimens before me, and is absolutely reliable. UPPER TENNESSEE DRAINAGE. 595 POWELL DRAINAGE. North Fork Powell River, Big Stone Gap, Wise Co., Va. Walker coll. (C. C. Adams, Sept. 4, 99). 1. Fusconaia pilaris bursa-pastoris 4. Eurynia nebulosa 2. F. barnesiana bigbyensis 5. Lampsilis fasciola 3. Medionidus plateolus South Fork Powell River, Big Stone Gap, Wise Co., Va. *— Walker coll. (Adams, Sept. 4, 99); t=—= Carnegie Mus. (Ortmann, May 15 and Sept. 6, ’13). +1. Fusconaia pilaris bursa-pastoris +7. Alasmidonta minor +2. F. barnesiana +8. Strophitus edentulus *+3, F. barnesiana Pree +9. Ellipsaria subtenta +4. Lexingtonia dolabelloides con- *10. Toxolasma lividum radi *+11. Medionidus plateolus *tc. Pleurobema oviforme argen- *f12. Eurynia nebulosa tewm +13. E. vanuxemensis +6. Lasmigona badia Powell River, Olinger, Lee Co., Va. Walker coll. (Adams, Sept. 5, ’99). 1. Fusconaia pilaris bursa-pastoris 7, Lasmigona costata 2. F. cuneolus 8. Alasmidonta marginata 3. F. barnesiana bigbyensts 9. Ellipsaria subtenta 4. Lexingtonia dolabelloides con- 10. Medionidus plateolus radi 11. Eurynia perpurpurea 5. Pleurobema oviforme 12. E. nebulosa 6. Elliptio dilatatus 13. Lampsilis ovata ventricosa Powell River, Dryden, Lee Co., Va. * — Walker coll. (Adams, Sept. 3, ’99) ; {=—=Carn. Mus. (Ort- mann, Sept. 7, 13). +1. Fusconaia pilaris bursa-pastoris +9. Alasmidonta marginata +2. F. barnesiana +10. Pegias fabula *+3. F. barnesiana bigbyensis +11. Strophitus edentulus *+4. Lexingtonia dolabelloides con- *f12. Ellipsaria subtenta radi *+13. Medionidus plateolus +5. Pleurobema oviforme *+14. Eurynia nebulosa *+6. Elliptio dilatatus *+15, EH. vanuxemensis +7. Lasmigona badia +16. Lampsilis ovata ventricosa *78. L. costata *+17. L. fasciola 596 ORTMANN—NAYADES OF Cane Creek, Pennington Gap, Lee Co., Va. Walker coll. 1. Fusconaia pilaris bursa-pastoris 2. Pleurobema oviforme Puckell Creek, Pennington Gap, Lee Co., Va. Walker coll. 1. Fusconaia cuneolus 3. Pleurobema oviforme 2. Lexingtonia dolabelloides con- radi Wallen Creek, Lee Co., Va. (near Jonesville). * — Walker coll.; }==Carn. Mus. (from G. H. Clapp). *r, Fusconaia barnesiana bigbyensis *4. Medionidus plateolus 72. Elliptio dilatatus 75. Eurynia nebulosa *3, Pegias fabula Powell River, Lytton Mill, Pennington Gap, Lee Co., Va. Walker coll. (Adams, Sept. 1, ’99). 1. Fusconaia pilaris bursa-pastoris 8. Lasmigona costata 2. F. cor analoga 9. Ellipsaria fasciolaris 3. F. barnesiana bigbyensis 10, Nephronaias pectorosa 4. Quadrula cylindrica strigillata 11. Euryma nebulosa 5. Lexingtonia dolabelloides con- 12. E. vanuxemensis radt 13. Lampsilis fasciola 6. Pleurobema oviforme 14. Truncilla haysiana 7. Elliptio dilatatus Powell River, Jonesville, Lee Co., Va. *=—= Walker coll. ;—=Carn, Mus: (Hartman coll:): ii *1, Fusconaia pilaris bursa-pastoris *4, Ellipsaria subtenta *2. Pleurobema oviforme *5. Toxolasma lividum *3. Elliptio niger *16. Lemiox rimosus Powell River, Rose Hill, Lee Co., Va. Walker coll. (Adams, Aug. 5, ’o1). 1. Fusconaia pilaris bursa-pastoris 8. Nephronaias pectorosa 2. F. cor analoga 9. Lemiox rimosus 3. F. barnesiana bigbyensis 10. Medionidus plateolus 4. Pleurobema oviforme 11. Eurynia nebulosa 5. Elhiptio dilatatus 12. Lampsilis fasciola 6. Alasmidonta marginata 13. Truncilla interrupta 7. Ellipsaria fasciolaris UPPER TENNESSEE DRAINAGE. 597 Powell River, Shawanee, Claiborne Co., Tenn. Walker coll. (Adams, Aug. 31, 99). CI ANAwWDHND H . Fusconaia pilaris bursa-pastoris ) _ F. pilaris lesweuriana { . F. cuneolus F. cor analoga F. barnesiana bigbyensis . Amblema plicata costata . Pleurobema oviforme . P. oviforme eee . Elliptio niger . E. dilatatus . Ellipsaria subtenta . Dromus dromas caperatus . Nephroaias ligamentina gibba Powell River, Bryant Shoals, Claiborne Walker coll. (Adams, Aug. 30, ’99). NAM BW ND H _ Fusconaia pilaris bursa-pastoris . F. cor analoga . Amblema plicata costata . Quadrula pustulosa . Quadrula cylindrica strigillata . Plethobasus cyphyus . Lexingtonia dolabelloides con- radt . Pleurobema oviforme } . P. oviforme argentewm\ 10. Bas Elliptio dilatatus Lasmigona costata 12. 13: Lan b& bw WN Ny ios) . Nephronaias pectorosa . Toxolasma lividum . Medionidus plateolus . Eurynia nebulosa . Lampsilis ovata . L. ovata ventricosa . L. fasciola . Truncilla triquetra . T. interrupta . T. haysiana _ T. torulosa gubernaculum . T. capseformis Co., Tenn. Alasmidonta marginata Ellipsaria fasciolaris . E. subtenta . Dromus dromas caperatus . Nephronaias ligamentina gibba . N. pectorosa . Lemiox rimosus . Medionidus plateolus . Eurynia nebulosa . Lampsilis fasciola . Truncilla interrupta . T. capseformis Powell River, Combs, Claiborne Co., Tenn. Carn. Mus. (Ortmann, Sept. 12, ’13 and Sept. 13, '15). I 2 3 4 5 6 7 8 9 10 II . see Fusconaia pilaris bursa-pastoris F. pilaris lesweuriana cuneolus F. cor analoga F. barnesiana bigbyensis Amblema plicata costata Quadrula cylindrica Plethobasus cyphyus Lexingtonia dolabelloides con- radt Pleurobema oviforme P. oviforme cane 12. 13% 14. Ti5e 16. 17. 18. 20. 10. 21. 22, 23. Elliptio dilatatus Lasmigona costata Alasmidonta marginata Strophitus edentulus Ellipsaria fasciolaris E. subtenta Dromus droma caperatus Nephronaias pectorosa Nephronaias ligamentina gibba Paraptera fragilis Proptera alata Lemiox rimosus 598 24. . Eurynia fabalis . E. nebulosa . E. vanuxemensis . E. recta 25 26 27 28 20. 30. ORTMANN—NAYADES OF Medionidus plateolus Lampsilis ovata L. ovata ventricosa 31. L. fasciola . Truncilla triquetra . T. interrupta . T. haysiana . T. lewist . T. torulosa gubernaculum . T. capseformis Powell River, Green's Ford, Umon Co., Tenn. Walker coll. (Adams, Aug. 23, ’99). (Adams says: Campbell Co., Walker: Claiborne Co.; the locality is, where the three counties meet, and the ford is, according to map —sheet Maynardville—rather in Union Oo AN AWNAW ND . Fusconaia pilaris bursa-pastoris . FE. pilaris lesueuriana . F. cuneolus F. barnesiana . Amblema plicata costata . Quadrula cylindrica . Elliptio niger . E. dilatatus . Lasmigona costata 10. Tits 12) 133 14. 15. 16. 17 18. Con)= Strophitus edentulus Ellipsaria fasciolaris E. subtenta Toxolasma lividum Medionidus plateolus Eurynia fabalis E, nebulosa Truncilla interrupta T. capseformis Powell River, Powell River P. O., Campbell Co., Tenn. Walker coll. (Adams, Aug. 23, ’99). WN H . Amblema plicata costata . Elliptio dilatatus . Lasmigona costata . Ellipsaria subtenta ie 6. 7. Eurymnia fabalis Truncilla triquetra T. capseformis CLINCH DRAINAGE. North Fork Clinch River, Tazewell, Tazewell Co., Va. Walker coll. (Adams). I . Alasmidonta minor 2. Eurynia nebulosa Clinch River, Tazewell, Tazewell Co., Va. Carn. Mus. (Ortmann, Sept. 19, 12). 1. Fusconaia barnesiana bigbyensis 2. Lasmigona badia UPPER TENNESSEE DRAINAGE. 599 Clinch River, Cedar Bluff, Tazewell Co., Va. Gari Mus, (Ortmann, Sept. 20, 12, and May 11, 713). hw NH . Fusconaia pilaris bursa-pastoris . F. barnesiana bigbyensis . Quadrula cylindrica strigillata . Lexingtonia dolabelloides con- radi . Pleurobema oviforme . P. oviforme ss Stee . Lasmigona badia 5 6 7. Elliptio dilatatus 8 9 . L. costata 10. Alasmidonta minor 11. Strophitus edentulus 12. Ellipsaria subtenta 13. Medionidus plateolus 14. Eurynia perpurpurea 15. E. nebulosa 16. Lampsilis ovata ventricosa 17. L. fasciola 18. Truncilla capseformis Clinch River, Richland, Tazewell Co., Va. Carn Mus: ((@rtmann, Sept. 20, *12))- . Fusconaia pilaris bursa-pastoris . FP, barnesiana . Quadrula cylindrica strigillata I 2 3. F. barnesiana bigbyensis 4 5 a \o ON . Lexingtonia dolabelloides con- radt . Pleurobema oviforme . P. oviforme Hone . Elliptio dilatus . Lasmigona badia 10. L. costata 11. Alasmidonta minor 12. A. marginata 13. Strophitus edentulus 14. Ellipsaria subtenta 15. Medionidus plateolus 16. Eurynia perpurpurea 17. E. nebulosa 18. Lampsilis ovata ventricosa 19. L. fasciola Clinch River, Raven, Tazewell Co., Va. Carn. Mus. (Ortmann, Sept. 21, ’12). mh WD N 5 6. 7 8 . Fusconaia pilaris bursa-pastoris . F. barnesiana bigbyensis . Quadrula cylindrica strigillata . Lexingtonia dolabelloides con- radi . Pleurobema oviforme —) P. oviforme argenteum\ . Elliptio dilatatus . Lasmigona costata 9. Strophitus edentulus 10. Ellipsaria subtenta 11. Medionidus plateolus 12. Eurynia perpurpurea 13. E. nebulosa 14. Lampsilis ovata ventricosa 15. L. fasciola 16. Truncilla haysiana 17. T. capseformis Clinch River, Cleveland, Russell Co., Va. *—- Walker coll. (Adams, Aug., ’99); {—Carn. Mus. (Ort- mann) May 13> 713)). le #9 Fusconaia pilaris bursa-pastoris . F. cor analoga +3. Amblema plicata costata *+4. Quadrula intermedia 600 T5- *+6 17. 78. 19. *+10. TT. oe), +13: f14. ORTMANN—NAYADES OF Q. cylindrica strigillata Lexingtonia dolabelloides con- radt Pleurobemaobliquum coccineum P. oviforme P. oviforme argenteum | Elliptio dilatatus Lastena lata Lasmigona costata Alasmidonta minor Alasmidonta marginata Strophitus edentulus Ellipsaria fasciolaris E. subtenta Nephronaias pectorosa Medionidus plateolus Eurynia fabalis E. perpurpurea E. nebulosa Lampsilis ovata ventricosa L. fasciola Truncilla capseformis TIS. *+16, eye *+78, *TTO. *+20, TOT ozs 123: *12A, *25. Clinch River, Fink, Russell Co., Va. Carn. Mus. (Ortmann, May 12, 13). ON An BRwhd H . Fusconaia pilaris bursa-pastoris . F. cor analoga . F. barnesiana bigbyensis Amblema plicata costata . Quadrula cylindrica strigillata Pleurobemaoviforme argenteum . Elliptio dilatatus . Lasmigona costata 9. Alasmidonta marginata 10. Ellipsaria fasciolaris 11. Nephronaias pectorosa 12. Medionidus plateolus 13. Eurynia nebulosa 14. Lampsilis ovata ventricosa 15. L. fasciola 16. Truncilla capseformis Clinch River, St. Paul, Wise Co., Va. *—= Walker coll. (Adams, Aug. 8, ’99); {==Carn. Mus. (Ort- mann, May 14, ’13). Tite T2y 13. ap T5- +6. *t7- 78. *79. TIO. a Tle Tle: Fusconaia pilaris bursa-pastoris F. cor analoga F. barnesiana bigbyensis Amblema plicata costata Lexingtonia dolabelloides con- radt Pleurobema oviforme argenteum Elliptio dilatatus Lastena lata Lasmigona costata Alasmidonta minor A. marginata Strophitus edentulus *+13. Ellipsaria fasciolaris *t14. Ellipsaria subtenta +15. Nephronaias ligamentina gibba *+16. N. pectorosa ; +17. Lemiox rimosus *118. Medionidus plateolus t19. Eurynia fabalis 20. E. perpurpurea *tat. EH. nebulosa $22. E. recta 423. Lampsilis ovata ventricosa *t24, L. fasciola *t25. Truncilla capseformis Clinch River, Dungannon, Scott Co., Va. Walker coll. (Adams, Aug. I1, ’99). I. Amblema plicata costata 2. Truncillatorulosagubernaculum UPPER TENNESSEE DRAINAGE. 601 Clinch River, Clinchport, Scott Co., Va. * —- Walker coll. (Adams, Aug., 99); {—=Carn. Mus. (Ort- Manin, eplys, 713). ST *T2: 3. *T4. 15. *+6, 7 78. *9, FIO. Tra ty Te qs. *TTA, anos *16, 7 Fusconaia pilaris bursa-pastoris F, pilaris lesueuriana ! F. cuneolus F, cor analoga F. barnesiana bigbyensis Amblema plicata costata Quadrula intermedia Q. cylindrica Rotundaria tuberculata Plethobasus cyphyus Lexingtonia dolabelloides con- radi Pleurobemaobliquum coccineum P. oviforme Elliptio niger E, dilatatus Lastena lata Lasmigona costata *+78. *TIO. 720. TD Te *+22, *723. *+24, +25. $20. $27. 728. *+20,. *730. 731. 132: #133. #134. Alasmidonta marginata Ellipsaria fasciolaris E, subtenta Nephronaias ligamentina gibba N. pectorosa Proptera alata Medionidus plateolus Eurynia perpurpurea E. nebulosa E. recta Lampstlis ovata L. ovata aneee L. fasciola Truncilla triquetra T. interrupta T. torulosa gubernaculum T. capseformis Clinch River, Speer’s Ferry, Scott Co., Va. Carn. Mus. (Ortmann, July 8, ’13). CON AM BW ND H (=) Ne) x Ss MH Ww NH . Fusconaia pilaris bursa-pastoris . F. cuneolus . F. cor analoga . Amblema plicata costata . Quadrula cylindrica . Rotundaria tuberculata . Pleurobema oviforme argenteum . Elliptio dilatatus . Lasmigona costata . Alasmidonta minor . A, marginata . Ellipsaria fasciolaris . Nephronaias pectorosa (seen) 14. Toxolasma lividum TGs 16. 17 = TS 10. 20 21. 22. PAS Medionidus plateolus Euryma fabalis . E. trabalis E. perpurpurea E. nebulosa . BE. vanuxemensis Lampsilis ovata ventricosa Truncilla triquetra . T. interrupta . I. lenior T. capseformis Clinch River, Church Ford, Scott.Co., Va. Walker coll. Tre Fusconaia cor analoga PROC. AMER. PHIL. SOC., VOL. LVII, U, OCT. I, 1918. 602 ORTMANN—NAYADES OF Clinch River, Horton Ford, Hancock Co., Tenn. Walker coll. (Adams, Aug. 14, ’99). 1. Fusconaia pilaris bursa-pastoris 2. F. cuneolus 3 4 . Rotundaria tuberculata . Lasmigona costata Be Nephronaias ligamentina gibba 6. N. pectorosa Ze 8. Truncilla triquetra T. capseformis Clinch River, Kyle Ford, Hancock Co., Tenn. Walker coll. mb WwW NH . Fusconaia pilaris bursa-pastoris . F. pilaris lesueuriana . F. barnesiana . Pleurobema oviforme . T. oviforme argenteum 6. Elliptio dilatatus 7. Ellipsaria subtenta 8. Medionidus plateolus 0. Truncillatorulosagubernaculum Clinch River, Sneedville, Hancock Co. Tenn. (“between Kyle Ford and Sneedville”). Walker coll. (Adams, Aug. 16, ’99). 1. Fusconaia pilaris bursa-pastoris 2 3 . F. cor analoga . Plethobasus cyphyus 4. 5. Elliptio dilatatus Ellipsaria subtenta Clinch River, Oakman, Grainger Co., Tenn. Carn. Mus) (Orimann, May 25715; Sept) t44015)): ON AM BRW ND A Ke) . Fusconaia pilaris bursa-pastoris . F. cuneolus . Amblema plicata costata . Rotundaria tuberculata . Plethobasus cyphyus . Pleurobema obliquum rubrum . Elliptio niger . E. dilatatus . Lastena lata . Lasmigona costata . Alasmidonta marginata . Strophitus edentulus . Ellipsaria fasciolaris . Cyprogenia stegaria . Nephronaias ligamentina gibba 16. 17. 18. 19. 20. Rite 22. 23% 24. QE 26. 27 28 20. 30. Nephronaias pectorosa Amygdalonaias truncata Proptera alata Medionidus plateolus Eurynia nebulosa E. vanuxemensis E. recta Lampsilis ovata L. ovata pee L. fasciola Truncilla triquetra T. arceformis T. interrupta T. torulosa gubernaculum T. capseformis UPPER TENNESSEE DRAINAGE. 603 Clinch River, Clinch River Station, Claiborne Co., Tenn. Canim Muss (Ortmann, Sept. 11;,753'): Oo ON An BW ND 10. Tale 12) Ts 14. 15. 16. 17: 18. . Cumberlandia monodonta . Fusconaia pilaris bursa-pastoris . F. cuneolus . F. cor analoga . Amblema plicata costata . Ouadrula cylindrica . Rotundaria tuberculata . Plethobasus cyphyus ; . Lexingtonia dolabelloides con- radt Pleurobema obliquum rubrum Elliptio niger E. dilatatus Lasmigona costata Alasmidonta marginata Strophitus edentulus Ellipsaria fasciolaris E, subtenta Cyprogenia stegaria 19. 20. 21 22. 23. 24. 25. 34. 35. 36. 37- Dromus dromas caperatus Nephronaias ligamentina gibba N. pectorosa Amygdalonatas truncata Paraptera fragilis Proptera alata Medionidus plateolus . Eurynia fabalis . E. nebulosa _ E récta . Lampsilis ovata . L. ovata ventricosa . L. fasciola . Truncilla triquetra . arceformis . interrupta . haysiana . torulosa gubernaculum . capseformis bop ate liga setae) Clinch River, Black Fox Ford, Union Co., Tenn. *— Walker coll. (Adams, Aug. 19, mann, Sept. 15, 715). TT, 2: #13: 14. Be 76. 17. #78. QO. a KO rte 12 13s *T 4, ads Fusconaia pilaris bursa-pastoris ) F, pilaris lesweuriana § F. cuneolus Amblema plicata costata Rotundaria tuberculata (seen) Plethobasus cyphyus Pleurobema obliquum catillus P. obliquum rubrum Elliptio dilatatus (seen) Lasmigona costata (seen) Alasmidonta marginata Strophitus edentulus (seen) Ellipsaria fasciolaris E. subtenta Cyprogenia stegaria F16 T17 *F18 19. +20. T2T. 22. 723: #24. $25. *126. 127. *F28, *+20, *7 30. 99) ; f==Carn. Mus. (Ort- . Nephronaias ligamentina gibba . N. pectorosa . Amygdalonaias truncata Proptera alata (seen) Medionidus plateolus Eurynia nebulosa E. recta (seen) Lampsilis ovata L. ovata ventricosa L. fasciola Truncilla triquetra T. interrupta T. haysiana T. torulosa gubernaculum T. capseformis Clinch River, Walker's Ford, Union Co., Tenn. Walker coll. I. Fusconaia barnesiana ° 604 ORTMANN—NAYADES OF Clinch River, Needham Ford, Union Co., Tenn. Walker coll. (Adams, Aug. 19, ’99). Ke) CON anbw ND He . Cumberlandia monodonta . Fusconaia pilaris bursa-pastoris . F, pilaris lesueuriana ENGOr, F. cor analoga . Rotundaria tuberculata . Plethobasus cyphyus . Pleurobema obliquum ) . P. obliquum coccineum | . P. obliquum rubrum . Elliptio niger . E. dilatatus . Lasmigona costata . Alasmidonta marginata . Strophitus edentulus . Ellipsaria fasciolaris . Cyprogenia stegaria . Dromus dromas caperatus . Nephronaias ligamentina gibba . N. pectorosa . Paraptera leptodon . Proptera alata . Eurynia recta 24. 25. 26. Lampsilis ovata L. fasciola Truncilla interrupta Clinch River, Kelly Ford, Union Co., Tenn. Walker coll. (Adams, Aug. 20, ’99) (“between Kelly and Sharp’s Ford”). Te Cumberlandia monodonta 2. Truncillatorulosagubernaculum Clinch River, below Agee, Campbell Co., Tenn. Walker coll. (Adams, Aug. 24, ’99) (“Agee to Offutt”). 1. Cumberlandia monodonta 2. Fusconaia pilaris lesueuriana 3. Quadrula cylindrica 4. Rotundaria tuberculata EB 6 7 8 Plethobasus cyphyus . Lexingtonia dolabelloides . Pleurobema obliquum . P. obliquum rubrum 0. 10. II 14 15 16 Elliptio niger E, dilatatus . Ellipsaria fasciolaris 12: 13s Obliquaria reflexa Cyprogenia stegaria . Dromus dromas caperatus . Nephronaias ligamentina gibba . Eurynia recta Clinch River, Offutt, Anderson Co., Tenn. Carn. Mus. (Ortmann) Sept. 1914). © CONT OV OW BS G Wb) ie -— = eH Oo HO . Cumberlandia monodonta . Fusconaia pilaris bursa-pastoris } . F, pilaris lesueuriana . F. cuneolus appressa . Amblema plicata costata . Quadrula pustulosa . Rotundaria tuberculata . Plethobasus cyphyus . Pleurobema obliquum . P. obliquum rubrum . Elliptio niger . E. dilatatus 1gy- 14. iis 16. 7 18. 19. 20. 2. 22 23 Lasmigona costata Ellipsaria fasciolaris Cyprogenia stegaria Dromus dromas caperatus Nephronaias ligamentina gibba - Plagiola lineolata : Proptera alata — Eurynia recta Lampsilis ovata . L. fasciola (seen) . L. orbiculata UPPER TENNESSEE DRAINAGE. 605 Clinch River, Clinton, Anderson Co., Tenn. (Moore Ferry). * —= Walker coll. (Adams, Aug. 25, ’99) (‘‘ between Offutt and Clinton”) ; f==Carn. Mus. (Ortmann, Sept. 7, 14) ; (no. 42: Hart- man coll.). I. *2, miss 14. ies *16. WA 78. 79. *+T0, toranilee F12. ales T14. FTTS. F106. eT 7. +18. *TTO. *+20, 2s 22) Cumberlandia monodonata (seen ) Fusconaia pilaris F. pilaris lesweurtana F. pilaris bursa-pastoris F. cuneolus F, cuneolus aul, F, barnesiana Amblema plicata costata Quadrula pustulosa O. cylindrica Rotundaria tuberculata Plethobasus cyphyus Lexingtonia dolabelloides L. dolabelloides conradi ! Pleurobema obliquum P. obliquum Sea P. obliquum rubrum P. oviforme Elliptio niger E. dilatatus Lastena lata Lasmigona costata (seen) *+23. Alasmidonta marginata 24. Strophitus edentulus *t25. Ellipsaria fasciolaris 726. E. subtenta 27. Obliquaria reflexa *+28. Cyprogenia stegaria +29. Dromus dromas caperatus +30. Obovaria retusa *+31. Nephronatas ligamentina gibba *+32, Amygdalonias truncata 33. Proptera alata (seen) 434. Lemiox rimosus 735. Eurynia vanuxemensis *+36. FE. recta *+37, Lampsilis ovata 738. L. ovata ventricosa +39. L. fasciola *+40. Truncilla triquetra +41. T. haysiana +42. T. stewardsoni (Hartm. coll.) +43. T. propinqua +44. T. capseformis Clinch River, Edgemoor, Anderson Co., Tenn. Carns Muss (Ortmann, Sept. 514) Sept, 17,715). Lon! HOOD ON ANB W ND i He NOW mn RW db . Cumberlandia monodonta . Fusconaia pilaris . F, pilaris lesueuriana F. pilaris bursa-pastoris | F. cor F., barnesiana F. barnesiana tumescens | . Ouadrula pustulosa . O. cylindrica . Rotundaria tuberculata . Plethobasus cooperianus . P. cyphyus . Pleurobema obliquum . P. abliquum Ein | P. obliquum catillus | 16. P. obliquum rubrum 17. P. oviforme holstonense 18. Elliptio niger 19. E. dilatatus 20. Lastena lata 21. Lasmigona costata 22. Alasmidonta marginata 23. Strophitus edentulus 24. Ellipsaria fasciolaris 25. Obliquaria reflexa 26. Cyprogenia stegaria 27. Dromus dromas caperatus 28. Nephronaias ligamentina gibba 29. Amygdalonias truncata 20. Paraptera leptodon Truncilla triquetra . T. capseformis 606 ORTMANN—NAYADES OF 31. P. fragilis 37. 32. Proptera alata 38. T. interrupta 33. Eurynia recta 39. T. haysiana 34. Lampsilis ovata 40. T. propinqua 35. L. ovata beeen 41 36. L. fasciola Clinch River, Solway, Knox Co., Tenn. Carn. Mus. (Ortmann, Sept. 12, ’14). 1. Cumberlandia monodonta 18. 2. Fusconaia pilaris 10. 3. F. pilaris lesueuriana 20. 4. F. pilaris bursa-pastoris 2% 5. F. cuneolus appressa 22} 6. F. barnesiana bigbyensis 22% 7. Amblema plicata costata 24. 8. Quadrula pustulosa 25. 9. Q. cylindrica (seen) 206. 10. Rotundaria tuberculata Py fe 11. Plethobasus cyphyus 28. 12. Lexingtonia dolabelloides 2 13. Pleurobema obliquum 30. 14. P. obliquum cordatum Bite 15. P. obliquum coccineum Sy 16. P. obliquum rubrum Gey 17. Elliptio niger 34. Clinch River, Patton’s Ferry, Roane Co (According to Pilsbry & Rhoads.) ON AND wW HD H . Fusconaia pilaris 0. F. cor 10. F. barnesiana tumescens Tile . Amblema plicata costata 12) . Quadrula pustulosa 1S: . Plethobasus cooperianus 14. . Pleurobema obliquum rubrum 15. . Elliptio niger 16. =, Elliptio dilatatus Strophitus edentulus Obliquaria reflexa Cyprogenia stegaria Dromus dromas caperatus Nephronaias ligamentina gibba Amygdalonaias truncata Paraptera fragilis (seen) Proptera alata Eurynia nebulosa E. recta . Lampsilis ovata L. ovata ventricosa L. orbiculata Truncilla triquetra T. haysiana T. capseformis Tenn, Elliptio dilatatus Alasmidonta marginata Obliquaria reflexa Nephronaias ligamentina gibba Plagiola lineolata Eurynia vanuxemensis Lampsilis fasciola Truncilla propinqua (SMALL TRIBUTARIES OF CLINCH.) Panther Creek, Hancock Co., Tenn. (near Sneedville, also spelled Paimters@reek): Walker coll. 1. Strophitus edentulus UPPER. TENNESSEE DRAINAGE. 607 Cove Creek, Caryville, Campbell Co., Tenn. *—=+ Walker coll.; +=—Carn. Mus, (Ortmann) Sept. 12) 15). y1. Fusconaia barnesiana +4. Medionidus plateolus +2. F. barnesiana bigbyensis +5. Eurynia nebulosa *+3, Lasmigona badia 76. E. vanuxemensis Coal Creek, Anderson Co., Tenn. “— Walker coll). 3——reported by Call: *tr, Fusconaia barnesiana *2, Lampsilis virescens Cane Creek, Offutt, Anderson Co., Tenn. (Ontmann,, Sept 414). 1. Eurynia vanuxemensis (seen) Bull Run, Heiskell, Knox Co., Tenn. Walker coll. 1. Lasmigona badia (DRAINAGE OF POPLAR CREEK.) Brush Fork, Marlow, Anderson Co., Tenn. Carn. Mus. (Ortmann, Sept. 2, ’14). 1. Alasmidonta minor 3. Eurynia vanuxemensis 2. Eurynia nebulosa Poplar Creek, Roane Co., Tenn. -—\\Valker colle; — reported by Call *1. Fusconaia cuneolus *4. Amblema plicata costata *t2,. F. cuneolus appressa *5. Pleurobema oviforme *32. F. cor Emory River, Harriman Junction, Roane Co., Tenn. Carn. Mus. (Ortmann, May-15, 715). 1. Elliptio niger 2. Eurynia nebulosa 608 ORTMANN—NAYADES OF Emory River, Harriman, Roane Co., Tenn. ~—according to Pilsbry & Rhoads; ,=Carn. Mus. (Ortmann, May 16, ’I5). ty1. Fusconaia cuneolus tr10. Toxolasma lividum to. F. barnesiana tumescens tir. Medionidus plateolus +3. Amblema plicata costata 12. Eurynia perpurpurea +4. Pleurobema oviforme 713. E. nebulosa ts. P. oviforme holstonense t14. E. vanuxemensis 6. Elliptio niger ~(seen) +15. Lampsilis virescens t}7. E. dilatatus 716. L. ovata ventricosa 8. Lasmigona costata (seen) 17. L. fasciola tto. Ellipsaria fasciolaris 718. Truncilla turgidula Houston DRAINAGE. (HEADWATERS. ) Little Mocassin Creek, Gate City, Scott Co., Va. Carn. Mus. (Ortmann, May 16, ’13). 1. Lasmigona badia 2. Eurymia vanuxemensis Big Mocassin Creek, Willow 2s Russell Co., Va. Walker coll. 1. Fusconaia barnesiana bigbyensis 2. Pleurobema oviforme argenteum Big Mocassin Creek, Mocassin Gap, Scott Co., Va. Carn! Mus. (Ortmann, May 16, ’13; Sept. 9, 715). 1. Fusconaia cuneolus 9. Ellipsaria subtenta 2. F. barnesiana 10. Medionidus plateolus 3. F. barnesiana Bee 11. Eurynia nebulosa 4. Ouadrula cylindrica 12. E. vanuxemensis 5. Pleurobemaoviforme argentewm 13. Lampsilis ovata ventricosa 6. Alasmidonta minor 14. L. fasciola 7. A. marginata 15. Truncilla capseformis 8 . Pegias fabula North Fork Holston River, Saltville, Smyth Co., Va. j= Carn. Mus. (Ortmann, Sept.17, “12)) 5) * = Carn, Muss (0; Avabeterson, jiile 20, -17 )c +*1. Fusconaia barnesiana bigbyensis +*3. Pleurobema oviforme argenteum +*2. Lexingtonia dolabelloides con- +4. Lasmigona costata radt +5. Alasmidonta minor 76. eae +8. 9. UPPER TENNESSEE DRAINAGE. 609 A, marginata Pegias fabula Strophitus edentulus Ellipsaria subtenta tio. Nephronaias pectorosa erate Toxolasma lividum *12. Medionidus plateolus 7*13. Eurynia nebulosa +*14. FE. vanuxemensis y15. Lampsilis ovata ventricosa 716. L. fasciola North Fork Holston River, Holston, Washington Co., Va. Walker coll. (Adams, Aug. 13, ’or). BW NH “TI. OQ U1 . Fusconaia cor analoga . FP, barnesiana . F. barnesiana bigbyensis . Lexingtonia dolabelloides con- radi . Pleurobema oviforme argenteum . Elliptio dilatatus . Lasmigona costata 8. Pegias fabula 9. Ellipsaria subtenta 10. Nephronaias pectorosa 11. Toxolasma lividum 12. Lemiox rimosus 13. Medionidus plateolus 14. Eurynia nebulosa 15. Lampsilis fasciola North Fork Holston River, Mendota, Washington Co., Va. *=— Walker coll: (Adams, Oct. 13, eo) ; {==Carn. Mus. (Ort tann, Jaly5,. <1). FIO. Gielen *+72, Tits . Fusconaia pilaris bursa-pastoris . F. cuneolus . F. cor analoga . F. barnesiana . F. barnesiana bigbyensis . Quadrula intermedia . O. cylindrica strigillata . Rotundaria tuberculata . Lexingtonia dolabelloides con- radt Pleurobema oviforme P. oviforme pee Elliptio dilatatus Lasmigona costata +14. Alasmidonta marginata 715. Pegias fabula 716. Ellipsaria fasciolaris *117. E. subtenta +18. Nephronaias pectorosa *19. Medionidus plateolus +20. Eurynia perpurpurea *t21, FE. nebulosa f22. E. vanuxemensis 723. Lampsilis ovata ventricosa 724. L. fasciola 425. Truncilla triquetra *26. T. capseformis North Fork Holston River, Hilton, Scott Co., Va. ‘Carn. Mus. (Ortmann, July 7, 13). Anup wd H . Fusconaia pilaris bursa-pastoris F. pilaris lesueuriana . FB. cuneolus . F. cor analoga . Amblema plicata costata . Quadrula cylindrica strigillata 7. Rotundaria tuberculata 8. Lexingtonia dolabelloides con- radi 9. Pleurobema oviforme ) 10. P. oviforme argenteum { 11. Elliptio dilatatus 12) 13. 14. 15. 16. 7a 18. 19. 20. ORTMANN—NAYADES OF Lasmigona costata Alasmidonta marginata Strophitus edentulus Ellipsaria fasciolaris Nephronaias pectorosa Lemiox rimosus Medionidus plateolus Eurynia fabalis E. perpurpurea 21. E. nebulosa 22. E. vanuxemensis 23. Lampsilis ovata ventricosa 24. L. fasciola 25. Truncilla triquetra 26. T. interrupta 27. T. haysiana 28. T. capseformis North Fork Holston River, Holston Bridge, Scott Co., Va. Walker coll. (Adams, Aug. II, ’or). ON Am RW ND 4 . Fusconaia pilaris lesueuriana . F. cor analoga . F. barnesiana bigbyensis . Quadrula cylindrica strigillata . Elliptio dilatatus . Lasmigona costata . Ellipsaria subtenta . Nephronaias ligamentina gibba 9. Nephronaias. pectorosa 10. Toxolasma lividum 11. Eurynia vanuxemensis 12. Lampsilis fasciola 13. Truncilla triquetra 14. T. interrupta 15. 7. torulosa gubernaculum 16. T. capseformis North Fork Holston River, Rotherwood, Hawkins Co., Tenn. Canna Muss (Ortmann, july 12, a3 Sepis5) 03): HOO ON Au KRW DH Lael H bo . Fusconaia pilaris Laie . pilaris lesueuriana . cuneolus . cor analoga . barnesiana . barnesiana ee 5) 35) 35) (25) 155) . Amblema plicata costata . Quadrula cylindrica strigillata . Rotundaria tuberculata . Plethobasus cyphyus . Lexingtonia dolabelloides con- radt . Pleurobema oviforme . 14. 15. 16. 17. 18. P. oviforme argenteum Elliptio dilatatus Lasmigona costata Alasmidonta marginata Strophitus edentulus Ellipsaria fasciolaris 19. Ellipsaria subtenta 20. Nephronaias ligamentina gibba 21. N. pectorosa 22. Proptera alata (seen) 23. Lemiox rimosus 24. Medionidus plateolus 25. Eurynia fabalis 26. E. perpurpurea 27. E. nebulosa 28. E. vanuxemensis 20. E: recta 30. Lampsilis ovata ventricosa 31. L. fasciola 32. Truncilla triquetra 33. T. interrupta 34. T. lentor 35. IT. haysiana 36. T. torulosa gubernaculum 37. T. capseformis Middle Fork Holston River, Marion, Smyth Co., Va. Carn. Mus. (Ortmann, Sept.. 16, 712). Te Eurynia nebulosa 2. Eurynia vanuxemensis UPPER TENNESSEE DRAINAGE: 611 Middle Fork Holston River, Chilhowie, Smyth Co., Va. Carn. Mus. (Ortmann, May 20, 13). 1. Fusconaia barnesiana-bigbyensis 2: 3; 4. 5. 6. Pleurobema oviforme argenteum Elliptio dilatatus Lasmigona costata Alasmidonta minor A. marginata ope 8. Q. 10. II. 12. Ellipsaria subtenta Medionidus plateolus Eurynia nebulosa E. vanuxemensis Lampsilis fasciola Truncilla walkert Laurel Creek, Mock’s Mill (near Vestal P. O.), Washington Co., Va, Walker coll. (Adams, Aug. 26, ’or). We Fusconaia barnesiana bigbyensis South Fork Holston River, Barron, Washington Co., Va. 7 — Walker coll=(Adams, Aue. 145or));*}— Carn, Muss (Ort- mann, May 19, ’13). pL. f2. #3 #4, is. *6, Fusconaia barnesiana bigbyensis Pleurobemaoviforme argenteum Elliptio dilatatus Lasmigona costata Alasmidonta minor A. marginata Ellipsaria subtenta +8: *19 *+1O *F II er, pias Nephronaias pectorosa Medionidus plateolus Eurynia nebulosa E. vanuxemensis Lampsilis fasciola Truncilla walkeri South Fork Holston River, Fish Dam, Sullivan Co., Tenn. Walker coll. (Adams, Sept. 6, ’or). Ti 2. ab Lexingtonia dolabelloides con- radt Elliptio dilatatus Nephronaias pectorosa 4. 5 6. Eurynia nebulosa E. vanuxemensis Lampsilis fasciola South Fork Holston River, Emmett, Sullivan Co., Tenn. Carn. Mus. (Ortmann, July 9, 713). 15 2. N ou & W& Fusconaia barnesiana bigbyensis Lexingtonia dolabelloides con- radi . Pleurobema oviforme } . P. oviforme argenteum { . Elliptio dilatatus . Lasmigona costata . Alasmidonta marginata 8. . Ellipsaria subtenta . Nephronaias pectorosa . Eurynia nebulosa . BE. vanuxemensis . Lampsilis ovata ventricosa . L. fasciola . Truncilla walkeri Ellipsaria fasciolaris 612 ORTMANN—NAYADES OF South Fork Holston, Bluff City, Sullivan Co., Tenn. Carn. Mus. (Ortmann, July 10, 13). Aum WN H Beaver Creek, Bristol, Washington Co., Carn. Mus. (Ortmann, July 6, ’13). He . Fusconaia pilaris bursa-pastoris | . F. pilaris lesueuriana . F, barnesiana F. barnesiana bibgyensis . Quadrula intermedia . Lexingtonia dolabelloides con- radt . Pleurobema oviforme argenteum . Elliptio dilatatus . Lasmigona badia Alasmidonta minor 10. . Alasmidonta marginata . Strophitus edentulus . Ellipsaria fasciolaris . E. subtenta (seen) . Nephronaias pectorosa . Eurynia nebulosa . E. vanuxemensis . Lampsilis ovata ventricosa . L. fasciola 2. Lasmigona costata Va. Medionidus plateolus South Fork Holston River, Pactolus, Sullivan Co., Tenn. Carn. Mus. (Ortmann, May 20, 14). OI AN RW DN A . Fusconaia pilaris bursa-pastoris ) . F. pilaris lesueuriana . F. barnesiana . Amblema plicata costata . Ouadrula intermedia . Q. cylindrica strigillata Rotundaria tuberculata . Pleurobema oviforme . P. oviforme argenteum . Elliptio niger . E. dilatatus . Lasmigona costata . Alasmidonta marginata . Pegias fabula . Strophitus edentulus . Ellipsaria fasciolaris . E. subtenta . Nephronaias ligamentina gibba . N. pectorosa . Medionidus plateolus . Eurynia fabalis . Lampsilis ovata ventricosa . L. fasciola . Truncilla triquetra . I. lemor . T. haysiana . T. torulosa gubernaculum . T. capseformis Watauga River, Watauga, Carter Co., Tenn. +—according to Pilsbry & Rhoads; }—=Carn. Mus. (Ortmann, HalyrrA,), 03). ale 72. +3. 14. Fusconaia pilaris bursa-pastoris F. barnesiana bigbyensis Pleurobema oviforme argenteum Lasmigona badia ts. +6. 7s +8. Nephronaias pectorosa Eurynia nebulosa E, vanuxemensis Lampsilis fasciola UPPER TENNESSEE DRAINAGE: 613 Watauga River, Johnson City, Washington Co., Tenn. According to Pilsbry & Rhoads. I . Fusconaia pilaris bursa-pastoris 2. Elliptio dilatatus 3} 4 5 . Lasmigona costata . Alasmidonta marginata . Strophitus edentulus 6. Medionidus plateolus 7. Eurynia nebulosa 8. E. vanuxemensis 9. Lampsilis fasciola (Hotston PROPER. ) Holston River, Church Hill, Hawkins Co., Tenn. Carne Must (Ortmann, Aue. 25, °14). I 2 Aum bh W Z. 8 . Fusconaia pilaris bursa-pastoris } . F, pilaris lesueuriana . Amblema plicata costata (seen) . Quadrula intermedia (seen) . O. cylindrica . Rotundaria tuberculata . Elliptio niger (seen) . E. dilatatus 9. Lasmigona badia 10. L. costata 11. Nephronaias ligamentina gibba . Proptera alata (seen) . Eurynia fabalis . Truncilla lenior I2 13 14 15 ae TON wae haysiana capseformis Holston River, Austin Mill, Hawkins Co., Tenn. *— Walker coll. (“ Rogersville,” practically the same locality) ; +=Carn. Mus. (Ortmann, Aug. 24, ’14). . Cumberlandia monodonta(seen) . Fusconaia pilaris lesweuriana (seen) . F, cuneolus . F. cuneolus appressa . F. barnesiana . Amblema plicata costata . Quadrula cylindrica . Rotundaria tuberculata . Pleurobema obliquum rubrum +10. pitts ee F13. Elliptio niger E. dilatatus Lasmigona costata Alasmidonta marginata F14. 715. 716. rT. *18, FIO. 720. 721. 122 *123, *2A, *F20,, *+26, Ellipsaria fasciolaris Nephronaias ligamentina gibba N. pectorosa Toxolasma lividum Lemiox rimosus Eurynia nebulosa Lampsilis ovata ventricosa Truncilla triquetra Te arceformis T. haysiana T. torulosa gubernaculum Ap T. capseformis turgidula 614 ORTMANN—NAYADES OF Holston River, Holston Station, Grainger Co., Tenn. *— Walker coll. (Adams); f==Carn. Mus. (Ortmann, Sept. Los LS). +1. Fusconaia pilaris lesueuriana *+20. Nephronaias pectorosa +2. F. cuneolus 721. Paraptera leptodon +3. F. barnesiana 722. P. fragilis *4. Rotundaria tuberculata (seen) +23. Proptera alata +5. Plethobatus cyphyus 724. Medionidus plateolus +6. Pleurobema obliquum 725. Eurynia recta +7. P. obliquum cordatum 726. Lampsilis ovata 78. P. obliquum rubrum $27. L. ovata eet to. P. oviforme 728. L. fasciola *10. Elliptio niger t29. Truncilla triquetra 711. E. dilatatus 730. T. arceformis 712. Lasmigona costata +31. T. interrupta +13. Alasmidonta marginata +32. T. haysiana +14. Strophitus edentulus +33. T. stewardsoni 715. Ellipsaria fasciolaris +34. T. lewisi 16. E. subtenta 735. T. turgidula *+17. Dromus dromas caperatus +36. T. florentina 718. Obovaria subrotunda levigata +37. T. capseformis y19. Nephronaias ligamentina gibba Holston River, Noeton, Grainger Co., Tenn. Carn. Mus. (Ortmann, May 22, ’14). 1. Cumberlandia monodonta 18. Lasmigona costata 2. Fusconaia pilaris 19. Strophitus edentulus 3. F. pilaris lesueuriana 20. Ellipsaria fasctolaris 4. F. cuneolus 21. E. subtenta 5. F. cuneolus renee 22. Dromus dromas caperatus 6. F. barnesiana 23. Nephronaias ligamentina gibba 7. F. barnesiana tumescens 24. Paraptera leptodon 8. Amblema plicata costata 25. P. fragilis 9. Quadrula cylindrica 26. Proptera alata 10. Rotundaria tuberculata 27. Eurynia fabalis 11. Pleurobema obliquum 28. E. nebulosa 12. P. obliquum catillus 29. E. recta 13. P. obliquum coccineum 30. Lampsilis ovata ventricosa 14. P. obliquum rubrum 31. Truncilla arceformis 15. P. oviforme 32. T. haysiana 16. Elliptio niger 33. T. turgidula cl 7. E. dilatatus 34. T. capsefornis UPPER TENNESSEE DRAINAGE. 615 Holston River, Turley Mull, Grainger Co., Tenn. Carn. Mus. (Ortmann, May 23, ’14). 1. Fusconaia pilaris 19. Nephronaias ligamentina gibba 2. F. pilaris lesueuriana 20. N. pectorosa 3. F. pilaris bursa-pastoris 21. Paraptera fragilis 4. F. cuneolus 22. Proptera alata 5. F. barnesiana tumescens 23. Lemiox rimosus 6. Amblema plicata costata 24. Eurynia nebulosa 7. Quadrula cylindrica 25. E. recta 8. Rotundaria tuberculata 26. Lampsilis ovata 9. Pleurobema obliquum rubrum 27. L. ovata ventricosa 10. P. oviforme 28. L. fasciola 11. Elliptio niger (seen) 29. Truncilla triquetra 12. E. dilatatus 30. T. arceformis 13. Lasmigona costata 31. T. interrupta 14. Alasmidonta marginata 2. T. haysiana 15. Strophitus edentulus 33. T. turgidula 16. Ellipsaria fasciolaris 34. T. florentina 17. Cyprogenia stegaria 35. IT. capseformis 18. Dromus dromas caperatus Holston River, McBee Ford, Hodges, Jefferson Co., Tenn. Carn. Mus. (Ortmann, May 25, ’14). 1. Cumberlandia monodonta(seen) 19. Strophitus edentulus 2. Fusconaia pilaris lesueuriana 20. Ellipsaria fasciolaris (seen) 3. F. barnesiana 21. E. subtenta (seen) 4. F. barnesiana tumescens 22. Cyprogenia stegaria 5. Amblema plicata costata 23. Dromus dromas caperatus 6. Quadrula pustulosa 24. Nephronaias ligamentina gibba 7. O. cylindrica 25. Amygdalonaias truncata 8. Rotundaria tuberculata 26. Paraptera fragilis 9. Plethobasus cyphyus 27. Proptera alata 10. Pleurobema obliquum 28. Eurynia recta 11. P. obliquum cordatum 29. Lampsilis ovata 12. P. obliquum coccineum 30. L. ovata ene 13. P. obliquum rubrum 31. Truncilla arceformis 14. P. oviforme 32. T. interrupta 15. Elliptio niger 33. T. haysiana 16. E. dilatatus 34. T. torulosa gubernaculum 17. Lasmigona costata 35. T. turgidula 18. Alasmidonta marginata 36. T. capseformis Holston River, Gant Island, near Strawberry Plains, Jefferson Co., Tenn. Walker coll. (Adams, Sept. 29, ’00). 1. Truncilla arceformis 616 ORTMANN—NAYADES OF Holston River, Mascot, Knox Co., Tenn. Carn. Mus. (Ortmann, Sept. 6, 714; Sept. 16, ’15). Ln oe | H Oo ON Aun i) b& bw b N HO bo wW HOO OWANDW DN H . Cumberlandia monodonta . Fusconaia pilaris . F, pilaris ee . F. barnesiana . Amblema plicata costata (seen) . Ouadrula pustulosa O. metanevra . QO. cylindrica . Rotundaria tuberculata . Plethobasus cyphyus . Pleurobema obliquum . P. obliquum cordatum ge P. obliquum rubrum . P. oviforme . P. oviforme holstonense . Elliptio niger . E. dilatatus . Lasmigona costata . Alasmidonta marginata . Strophitus edentulus . Ellipsaria fasciolaris . E. subtenta . Cyprogenia stegaria 44. 45. 46. . Dromus dromas .caperatus . Obovaria surotunda . O. subrotunda levigata . Nephronaias ligamentina gibba . N. pectorosa (seen) . Paraptera fragilis . Proptera alata . Lemiox rimosus . Medionidus plateolus . Eurynia fabalis . E. recta . Lampsilis ovata . L. ovata BA . L. fasciola . Truncilla triquetra . arceformis . interrupta . haysiana . stewardsoni . torulosa gubernaculum . turgidula . florentina . capseformis Ei aed eles Holston River, McMillan, Knox Co., Tenn. Garn. Mus. (Ortmann, Sept: 16, a3) . Cumberlandia monodonta . Fusconaia pilaris lesueuriana . Quadrula pustulosa QO. cylindrica . Rotundaria tuberculata . Plethobasus cyphyus . Pleurobema obliquum rubrum . Elliptio niger . E. dilatatus . Strophitus edentulus . Dromus dromas caperatus . Nephronaias ligamentina gibba . N. pectorosa 14. I5. 16. 17. 18. 10. 20. 2I =< Proptera alata Eurynia nebulosa E. recta Lampsilis ovata L. ovata ventricosa L. fasciola ° Truncilla triquetra . T. arceformis 22. 23% 24. 25. T. interrupta T. stewardsom T. florentina T. capseformis UPPER TENNESSEE DRAINAGE. 617 Holston River, Boyd Island, Knoxville, Knox Co., Tenn. According to Pilsbry & Rhoads (“1 mile above junction with French Broad’’). 1. Fusconaia pilaris 10. Cyprogema stegaria 2. Quadrula pustulosa 11. Dromus dromas 3. Q. cylindrica 12. Nephronaias ligamentina gibba 4. Rotundaria tuberculata 13. Paraptera fragilis 5. Pleurobema obliquum 14..Eurynia recta 6. P. obliquum rubrum 15. Lampsilis ovata 7. P. oviforme holstonense 16. Truncilla arceformis 8. Elliptio niger 17. T. propinqua g. E. dilatatus (SMALL TRIBUTARIES OF HOLSTON PROPER.) Big Creek, Rogersville, Hawkins Co., Tenn. Walker coll. 1. Lasmigona badia . 3. Medionidus plateolus 2. Alamidonta minor Big Flat Creek, Corryton, Knox Co., Tenn. Carn. Mus. (Ortmann, May 12, 14). 1. Fusconaia barnesiana 3. Eurynia Se loch 2. Lasmigona badia 4. E. vanuxemensis (seen) NoticHucky River DRAINAGE. Whitehorn Creek, Bulls Gap, Hawkins Co., Tenn. Carn. Mus. (Ortmann, May 18, ’14). 1. Lasmigona badia 3. Eurynia vanuxemensis 2. Eurynia nebulosa Bent Creek, Whitesburg, Hamblen Co., Tenn. Carn. Mus. (Ortmann, Sept. 8, ’15). 1. Lasmigona badia 3. Eurymia vanuxemensis 2. Eurynia nebulosa (seen) 618 ORTMANN—NAYADES OF Nolichucky River, Chunn’s Shoals, Hamblen Co., Tenn. Carn. Mus. (Ortmann, Sept. 17, ’13) (within a mile from con- fluence with French Broad). 1. Fusconaia cuneolus appressa 8. Paraptera fragilis 2. Amblema plicata costata 9. Proptera alata 3. Quadrula pustulosa 10. Eurynia recta 4. Rotundaria tuberculata 11. Lampsilis ovata ventricosa 5. Elliptio niger 12. Truncilla triquetra 6. Nephronaias ligamentina gibba 13. T. torulosa gubernaculum 7. N. pectorosa 14. T. capseformis FRENCH BROAD RIVER DRAINAGE. French Broad River, Asheville, Buncombe Co., N. Car. * — Walker coll. (Mrs. Andrews) ; {reported by Call. The French Broad at Asheville is now polluted by lumber indus- tries on Davidson River; repeated attempts on my part (May 13, A, cept. 14, 714, May 10, %15)) tailed to revealany *signs. of, the presence of shells. *1. Pleurobema oviforme (ravene- t4. Ellipsaria fasciolaris lianum) (also Lea) *t5. Medionidus plateolus *2. Elliptio dilatatus *t6. Eurynia nebulosa *3, Alasmidonta raveneliana (also *t7. Lampsilis fasciola Lea) £8. Truncilla capseformis Call also gives: Fusconaia pilaris and F. pilaris lesueuriana, but I have seen no specimens of these from this locality, and these records appear to me as very doubtful. French Broad River, Hot Springs, Madison Co., N. Car. According to Conrad, from ‘“‘ Warm Springs.” 1. Toxolasma lividum French Broad River, Boyd Creek, Sevier Co., Tenn. Carn. Mus. (Ortmann, Sept. 10, ’14). 1. Fusconaia pilaris 8. Plethobasus cooperianus 2. F, pilaris ee 9. P. cyphyus 3. F. barnesiana 10. P. cyphyus compertus 4. F. barnesiana een 11. Lexingtonia dolabelloides 5. Amblema plicata costata 12. Pleurobema obliquum i] 6. Quadrula pustulosa 13. P. obliquum cordatum { 7. Rotundaria tuberculata 14. Pleurobema obliquum rubrum UPPER TENNESSEE DRAINAGE. 619 15. P. oviforme 20. Ellipsaria fasciolaris 16. P. oviforme eet 21. Nephronaias ligamentina gibba 17. Elliptio niger 22. Proptera alata 18. E. dilatatus 23. Eurynia recta 19. Strophitus edentulus 24. Truncilla arceformis (TRIBUTARIES OF FRENCH BROAD.) Long Creek, near Bridgeport, Cocke Co., Tenn. + — Walker colli: —=reported: by Lea. *t, Lasmigona badia i t2. Eurynia vanuxemensis Big Pigeon River, Canton, Haywood Co., N. Car. Carn. Mus. (Ortmann, May 14, 714). 1. Alasmidonta raveneliana 2. Lampsilis fasciola Little Pigeon River, Sevierville, Sevier Co., Tenn. Carn. Mus. (Ortmann, Aug. 31, ’14). 1. Fusconaia barnesiana 8. Toxolasma lividum 2. F. barnesiana alent 9. Eurynia nebulosa 3. Pleurobema oviforme 10. E. vanuxemensis 4. P. oviforme peaelaeey 11. Lampsilis ovata ventricosa 5. Elliptio dilatatus 12) £. fasctola 6. Lasmigona badia 13. Truncilla capseformis 7. Alasmidonta minor Boyd Creek, Boyd Creek, Sevier Co., Tenn. Carn. Mus. (Ortmann, Sept. 5, ’14). 1. Fusconaia barnesiana 5. Lasmigona costata 2. Amblema plicata costata 6. Alasmidonta minor 3. Rotundaria tuberculata 7. Eurynia vanuxemensis 4. Elliptio dilatatus ; Pond, draining to French Broad, eight miles above Knoxville, Knox Co., Tenn. Walker coll (M. 1D: Barber): 1. Anodonta grandis gigantea 620 ORTMANN—NAYADES OF TRIBUTARIES OF TENNESSEE, BELOW KNOXVILLE. (FRoM THE WEST.) First Creek, Knoxville, Knox Co., Tenn. Walker coll. 1. Lasmigona badia Second Creek, Knoxville, Knox Co., Tenn. According to Lea. 1. Fusconaia barnesiana bigbyensis Third Creek, Knoxville, Knox Co., Tenn. Walker coll. 1. Lasmigona badia Piney River, Spring City, Rhea Co., Tenn. Carn. Mus. (Ortmann, May 18, ’15). 1. Lasmigona badia 2. Eurynia vanuxemensis Martin Branch of Sale Creek, Rhea Co., Tenn. Walker coll. (W. A. Marsh). (There is no: “ Martin Branch ” on map; Sale Creek flows from near Dayton, Rhea Co., south into Hamilton Co., and to the Tennessee. ) 1. Fusconaia barnesiana 2. F. barnesiana bigbyensis (FRoM THE East, COMING FROM THE MOUNTAINS.) Little River Drainage. Little River, Walland, Blount Co., Tenn. Carn. Mus. (Ortmann, May 12, ’15). 1. Fusconaia barnesiana bigbyensis 4. Euryma nebulosa 2. Pleurobemaoviforme argenteum 5. E. vanuxemensis 3. Medionidus plateolus UPPER TENNESSEE DRAINAGE. Little River, Melrose, Blount Co., Tenn. Carn. Mus. (Ortmann, Sept. 11, 715). 1. Fusconaia barnesiana bigbyensis 6. Eurynia nebulosa 2. Pleurobemaoviforme argenteum 7. E, vanuxemensis 3. Alasmidonta minor 8. Lampsilis ovata ventricosa 4. Nephronaias pectorosa 9. L. fasciola 5. Medionidus plateolus Little River, Rockford, Blount Co., Tenn. Carn. Mus. (Ortmann, Sept. 4, ’14). 1. Eurynia nebulosa Little River, Knox Co., Tenn. (lower part). + —— Walkers collts.p—= Carn. Mus: (Smith coll); *1, Cumberlandia monodonta *3. Truncilla triquetra +2. Amblema plicata costata Pistol Creek, Rockford, Blount Co., Tenn. Carn. Mus. (Ortmann, Sept. 4, ’14). 1. Fusconaia cuneolus appressa 4. Toxolasma lividum 2. F. barnesiana 5. Eurynia nebulosa 3. Alasmidonta minor LITTLE TENNESSEE RIVER DRAINAGE. Abram Creek, Blount Co., Tenn. Walker coll. 1. Fusconata barnesiana bigbyensis Tellico River, Monroe Co., Tenn. Walker coll., also reported by Lea. 1. Fusconaia barnesiana 2. F. barnesiana bigbyensis Little Tennessee River, Monroe Co., Tenn. * — Walker coll.; £— according to Marsh. tr, Fusconaia pilaris *5. Elliptio dilatatus *2, F. barnesiana tumescens *6, Lampsilis fasciola *3, Pleurobema oviforme *4. P. oviforme ieee 621 622 ORTMANN—NAYADES OF Little Tennessee River, Coytee, Loudon Co., Tenn. Walker coll. (Adams). 1. Fusconata pilaris 3. Truncilla haysiana 2. Lampsilis ovata 4. T. capseformis HIWASSEE RIVER DRAINAGE. Spring Creek, Austral, Polk Co., Tenn. Carn. Mus. (Ortmann, May 23, ’15). 1. Fusconaia barnesiana bigbyensis 3. Eurynia vanuxemensis 2. Eurynia nebulosa Cane Creek, McMinn Co., Tenn. Walker coll. (“ Cane Creek, Monroe Co.;” this is near Etowah, McMinn Co.). 1. Fusconaia barnesiana 3. Pleurobema oviforme 2. F. barnesiana bigbyensis Conasauga Creek, Monroe Co., Tenn. * — Walker coll.; £— according to Lewis. *1. Pleurobema oviforme *3, Lasmigona badia *t2. P. oviforme argenteum (also 4 *t4. Alasmidonta minor Lea) J t5. Eurynia nebulosa Ocoee River, Ducktown, Polk Co., Tenn. Walker coll. 1. Alasmidonta marginata Hiwassee River, Austral, Polk Co., Tenn. ‘Carn Mus. (Ortmann, Sept. 19, ’15). 1. Fusconaia barnesiana 5. Lasmigona badia 2. F. barnesiana tumescens 6. Eurynia trabalis 3. Pleurobema oviforme 4. P. oviforme holstonense Hiwassee River, Kincannon Ferry, Meigs Co., Tenn. Walker coll. (Adams, Oct. 9, ’or). 1. Elliptio niger UPPER TENNESSEE DRAINAGE. 623 Hiwassee River, Meigs Co., Tenn. Walker coll. (Adams). 1. Quadrula verrucosa 2. Lampsilis fasciola South Chickamauga Creek, Ringgold, Catoosa Co., Ga. + ——Walkenrcolla-(Adams, ‘Oct. 21, or)! 7—= Carn, Mus. ((Ort= mann, May 20, ’15). *t1. Pleurobema oviforme argenteum 78. Medionidus plateolus j2. Elliptio dilatatus 79. Eurynia trabalis +3. Lasmigona badia tio. E. nebulosa +4. L. costata 11. Lampsilis ovata ventricosa 75. Alasmidonta minor (seen) 76. Ellipsaria fasciolaris f12. L. fasciola 7. Obovaria subrotunda levigata TENNESSEE PROPER, AT AND BELOW KNOXVILLE. Tennessee River, at and above Knoxville, Knox Co., Tenn. ~—according to Pilsbry & Rhoads; ¢==Carn. Mus. (Ortmann, Septs22, 55 5. 4.—prabsonm Merry, and D. 1,— Dickerson Island), ti. Fusconaia pilaris +13. Dromus dromas to. F. barnesiana tumescens $14. Obovaria subrotunda +3. Quadrula pustulosa (B. F.) ty15. Nephronaias ligamentina gibba $4. Rotundaria tuberculata (DET) tt5. Plethobasus cooperianus (B. F.) $16. N. pectorosa +6. Pleurobema obliquum (B. F.) +17. Plagiola lineolata (B. F.) +7. P. obliquum rubrum t718. Proptera alata (B. F.) +8. P. oviforme holstonense t19. Lampsilis ovata tto. Elliptio niger (D. I.) t20. Truncilla triquetra tio. E. dilatatus tar. T. haysiana tir. Lasmigona costata t22. T. propinqua t12. Ellipsaria subtenta Tennessee River, three miles below Knoxville, Knox Co., Ten. Carn. Mus, (Ortmann, Sept: 21, 15): 1. Fusconaia pilaris 8. Pleurobema obliquum rubrum 2. F. pilaris ee 9. Elliptio niger 3. Quadrula pustulosa 10. E. dilatatus 4. Q. metanevra 11. Dromus dromas 5. Rotundaria tuberculata 12. Nephronaias ligamentina gibba 6. Plethobasus cooperianus 13. Plagiola lineolata 7. Pleurobema obliquum 14. Eurynia recta 624 ORTMANN—NAYADES OF Tennessee River, Little River Shoals, Knox Co., Tenn. * — Walker coll.; {according to Lewis. *1, Fusconaia barnesiana tumescens *6, Lasmugona costata *2. Pleuwrobema obliquum *7, Nephronaias ligamentina gibba *3, P. oviforme *8. Eurynia fabalis *4. P. oviforme holstonense to. Truncilla lewisi *5. Elliptio dilatatus Tennessee River, Concord, Knox Co., Tenn. + = according to Lewis; {== Carn. Mus. (Ortmann, Sept. 9, ’14). 71. Quadrula pustulosa 74. Elliptio niger {2. Rotundaria tuberculata +5. Strophitus edentulus £3. Pleurobema oviforme holsto- +6. Nephronaias ligamentina gibba nense +7. Eurynia recta Tennessee River, Chota Shoals, Knox Co., Tenn. Walker coll. 1. Fusconaia cuneolus appressa 3. Pleurobema oviforme holsto- 2. F. barnesiana nense Tennessee River (or “ Holston River”), Knox Co., Tenn. (Summary of all species reported from the region between Brab- son Ferry and Chota Shoals, mostly without exact locality.) ~= according to Lewis; §= reported by Pilsbry & Rhoads; += Walker coll.; {= Carn: Mus..((Hartman, coll? Ga.),/and Snirth coll. (S.)) ; those reported above are marked: B.= Brabson Ferry; 1D. Dickerson Island: K. == Knoxville: L.=Little, River Shoals; Co!—= Concord ; Ch. = Chota) Shoals: ti. Cumberlandia monodonta t12. Q. intermedia §+2. Fusconaia pilaris—K. +13. Q. cylindrica +73. F. pilaris lesueuriana—K., §t+14. Rotundaria tuberculata—K. Co. +4. F. cuneolus (?) t§t15. Plethobasus cooperianus—B. K. *5. F. cuneolus appressa—Ch. 16. P. cyphyus £6. F. cor $17. P. cyphyus compertus (also +*7. F. barnesiana—Ch. Frierson) t§*8. F. barnesianatumescens—K. L. +18. Lexingtonia dolabelloides to. Amblema plicata costata *t19. Pleurobema obliquum—B.K.L. ty10. Quadrula pustulosa—B. K. Co. t20. P. oliquum cordatum ty11. OQ. metanevra—K. t$f21. P. obliquum rubrum—K. UPPER TENNESSEE DRAINAGE. 625 t*22. P. oviforme—L. £§*23. P. oviforme holstonense—K. i. Con Eh. t§+24. Elliptio niger—D. K. Co. £§*}25.F. dilatatus—K. L. +26. Lastena lata £§*27. Lasmigona costata—K. L. $28. Alasmidonta marginata t}29. Strophitus edentulus—Co. $30. Ellipsaria fasciolaris £831. E. subtenta—K. +32. Obliquaria reflexa 4733. Cyprogenia stegaria (H.) £§+34. Dromus dromas—K. +35. Dromus dromas caperatus *+36. Obovaria retusa (S.)—K. £837. O. subrotunda—K. £8*+33. Nephronaias ligamentina gibba—D. K. L. Co. +830. N. pectorosa—K. +40. Amygdalonaias truncata tt41. Plagiola lineolata—B. K. +42. Paraptera leptodon 43. P. fragilis §+44. Proptera alata—B. K. 45. Toxolasma lividum 446. Lemiox rimosus 47. Medionidus plateolus 48. Eurynia fabalis—L, t49. E. nebulosa tso. E. vanuxemensis ty51. E. recta—K. Co. t§52. Lampsilis ovata—K. $53. L. fasciola *54. L. orbiculata £855. Truncilla triquetra—kK. $56. T. arceformis t57. T. interrupta *58. T. lenior t§s50. T. haysiana—K. t}60. T. stewardsoni (H.) (S.) 61. T. lewisi—L. t8}62. T. propinqua (S.)—K. +763. T. torulosa (H.)—K. 764. T. capseformis (H.) Anodontoides ferussacianus, reported by Lewis (as Anodonta oblita), has probably been misidentified. Also Fusconaia cuneolus is doubtfully given by Lewis; however, it may occasionally turn up in this region. Tennessee River, Hiwassee Island, Dayton, Rhea Co., Tenn. Walker coll. (Adams, Oct. 13, ’o1). 1. Amblema plicata costata 2. Pleurobema obliquum 3. Elliptio niger 4. Elliptio dilatatus 5. Ellipsaria fasciolaris Tennessee River, Rathburn, Hamilton Co., Tenn. Walker coll. (Adams). 1. Quadrula pustulosa . Rotundaria tuberculata . Lexingtonia dolabelloides . Pleurobema obliquum cordatum . P. obliquum rubrum . P. oviforme holstonense Am kw hd 7. Elliptio niger 8. E. dilatatus 9. Obliquaria reflexa 10. Cyprogenia stegaria 11. Dromus dromas 12. Plagiola lineolata 626 ORTMANN—NAYADES OF UPPER TENNESSEE DRAINAGE. Tennessee River, Chattanooga, Hamilton Co., Tenn. *=—according to Pilsbry & Rhoads; *== Walker coll. (Adams, Oct. 22,007 p= Carn. Mus: (juny coll) i 2 $3 $4 t5 +6 . Quadrula pustulosa . QO. metanevra . Plethobasus cooperianus . Pleurobema obliquum . P. oviforme holstonense . Elliptio niger 47. Elliptio dilatatus 8. Ellipsaria fasciolaris t*o. Dromus dromas t10. Nephronatas ligamentina gibba fir. Truncilla torulosa ii ee i, io ) pan . 4 i : 7 u - = _ : ‘tae een. Sha 7 « S few ¥ ) IADES kbar NA USSELS ( By A. E, ORTMANN, Pa. SoclETY PHICAL oso printed from — eS THE AMERICAN PHIL I NGs oe oe “Ri l@Erie Waterford Lorubee Worre, . . Edinboro COLLEC TING STAI IONS 4 Cam bridge Sp. 1 = 1 gy Meodville Hickory i as “\Cochrant Tionesta® IN THE DRAINAGES en i Molnet Bena eal mestown Ulicg A Sy henangy glerion 5 zy OF THE Shorr villa - oe alt \é, ok Leovittsburg | IGE, ged 8 Newton Fils Pulds ki wont O H I O bad fie jens Neshannack Fi/s: 44 astbrook empleton ‘ein Over lahoningtown Rosston Wompees Johnetta oy Creekside TEN N [ado Godfrey Aladdin —@South Bend ES S E E R IVE R S “\ Conners a Sheffield alrdga aon en (Gore polis Pittsburgh C A oY eaten izabeth gg “ss SCALE ' Ch oO 25 50 7s Wheeling i daity MILES A mune MMi lisboro xs Ten iley co : Mt. Morrj Pt. Marion | ain Doe es Cheat Ha’ anu ie oe aa ad Six eae | Marietta Y . ' tt Marys i I & ch Mine ' Cincinnati Parkersburg 5 " ! ornwallis 3f,. ! Hughes Lightburn ! SS rantsville WES SF = ‘Qnawha Burnasvil a) iF Pycoks Cossawaygedutton A ing Ramo d City owe la Fe © y Ue Shelton We af E sXharleston Y, On, np @ f°) ne <, Gq SiN wo °a7 ~ KJ 3 a ay GS a = eo Prestons burr - 4 - o Wem ~Tax€éwell 7 ~ a Cumberlan oH y x: 1 Nighlan OF ws Big Stop P- o~ SalivjHess ry eh. Olinger hilthowie VorEwilteg por Holston BONS rs RoseAill @ Speers Fy\ Moc&G, cae Avthore Batroree ah Re Hi Barron GC K ’ iorton Fd, Gotherwood hD. Se See ea ew fees BEE SGG, Clinch R.S. Seed Chufth Hire€actolus se aay eer Walker Fa @ydakina 4 nce A Mill Em mett; lack OC Ra uate Bluff City Agee €e Needham Fa. 0 7olston St. Wa 4 9P— = Aston e on ca 5 Offett nto cco ay : Turley Mill oa Shen.=Shenango = Mah. = = Mahoning Nesh. = Neshonnock Clinton YaHodges yr-rc7 Rac.= Raccoon Chart. >Chartiers Edgemoor Millan / Te M = 1K Mil [ Solway Kinoxwille ec =—— SUIS UC US Dunk. = Dunkard = Moc. Cr. = Mocassin Creek Mle RSROMWACr. WB aan rg Chotash' ¢BSevier Wile Southwestern continuation of map at A-B A Coy te ‘ GA + Asheville owe? oy BEE : Monroe Ver e Co. j Nkssec AustrajJ/~~—} ! San drainage ae eer ee - - Iroinage CORRELFATION OF SHAPE AND STATION IN FRESH- WATER MUSSELS (NATADES): By A. E. ORTMANN, Pu.D. (Read April 24, 1920.) While studying the Naiad-shells of the upper Ohio-drainage, the fact was forced upon my mind, that certain species which inhabit the headwaters and smaller streams are represented, in the larger streams, by different, but very similar forms, which are distin- guished from them chiefly by one character, namely obesity. The headwaters-forms are rather compressed or flat, the large-river- forms more convex and swollen. I also found that in the rivers of medium size integrades betwen the extremes were actually present. Subsequently, similar conditions appeared to prevail elsewhere, and the existence of this rule was brought home to me very forcibly during my study of the Naiad-fauna of the upper Tennessee drainage. Also other authors have observed this fact, and have referred to it in their publications. Thus Wilson and Clark’ indicated it for certain species of the Cumberland River (Amblema peruviana and costata; Rotundaria tuberculata and granifera; Dromus dromas and caperatus) and Utterback? has found this law to hold good in Missouri (chiefly in the Osage River) for several species (Fus- conaia undata and trigona; Amblema peruviana and costata; Pleuro- bema obliquum and allied forms). I myself have alluded to this relation between a compressed headwaters-form and a swollen large-river-form in the case of Obovaria lens and subrotunda,? in the case of Pleurobema coccineum 1 Bur. Fisher. Doc. no. 781, 1914, pp. 21 and 63. 2 Amer. Midl. Natural., 4, 1916, p. 2. 3 Ann. Carn. Mus., 5, 1900, p. 192. Reprinted from Proceedings American Philosophical Society, Vol. lix., 1920. 270 ORTMANN—CORRELATION OF SHAPE AND _and obliquum* and in the case of the group of Fusconaia barnesiana.® I have given a fuller account, including the synonymy of the forms concerned. Additional references may be found in my paper on the upper Tennessee shells® and on the Pennsylvanian Naiades.’ However, all this information is rather vague, and not supported by detailed measurements, and, before we finally assume that this law exists at all, we should substantiate it by more careful investi- gations. The present paper is written with the purpose to supply the details and to reduce them to figures. When we speak of the obesity of mussel shells, and call them flat and compressed, or convex and swollen, we refer to the pro- portional diameter of the shell. This diameter can be expressed in various ways, but I found that the most easy is in percentage of the length. The length is always measured parallel to the liga- ment,* which can be done, without difficulty, by placing the shell between the arms of a vernier caliper measure in the proper posi- tion. The diameter is measured at right angles to it, and always the maximum diameter of the two valves is taken, no matter where it is located. The length is then taken as=100, and the diameter ex- pressed in hundredths of the length. The material studied has been collected, for the largest part, by myself, and thus I am able to vouch for the correctness of the localities. It forms part of the collections of the Carnegie Museum in Pittsburgh. Yet there is additional material from other sources. The Carnegie Museum possesses a fine collection from the Tennessee drainage in northern Alabama and southern Tennessee made by H. 4 Nautil., 23, 1910, p. 117, footnote 2, and Ann. Carnegie Mus., 8, 1912, p. 264. 5 Nautil., 31, 1917, p. 58. 9 Proc. Amer. Philos. Soc., 57, 1918, p. 521 ff. “Mem. Carnegie Mus., 8, 1919. 8 The greatest length of the shell is sometimes in a diagonal direction, when the shell is “ oblique”; but since the obliquity varies much individually, the length parallel to the ligament is selected. Previous authors (Lea for instance) have not strictly adhered to this rule, and thus we may explain certain discrepancies of the figures given in his text, and those taken from his illustrations. In my measurements, I have always taken Lea’s illustra- tions as the standard. STATION IN FRESH-WATER MUSSELS. 271 i H. Smith; and finally I had the privilege of examining upper Ten- nessee shells in the collection of Mr. B. Walker in Detroit, col- lected by C. C. Adams (particulars about these may be found in my paper on the upper Tennessee Naiades). Of course, the shells collected at one and the same locality vary somewhat, and thus it was necessary to compute the average for each set. In some cases, only few specimens were at hand, in others, a great many of them. Thus I do not give, in the tables, the individual measurements, but only the extremes and the average. We shall see that this is entirely sufficient to establish the law. In each river, the localities are arranged from downstream upward, taking up the tributaries of the main stream generally in ascending order. The exact situation of each locality may be found on the map (page 258). According to obesity previous authors frequently have dis- tinguished separate species, and often corresponding forms in dif- ferent river-systems have received different specific names. Thus it has been a difficult task to bring order in the nomenclature, and I am compelled to devote considerable space to this. Since it will become apparent that the various groups of forms treated actually are connected by intergrades, each group should be regarded, taxo- nomically, as one species. But since, on the other hand, the ex- tremes are often rather sharply contrasted, and since they very generally appear as geographical (or rather ecological) races, it is convenient and justified, to use the old specific names in a varietal (or subspecific) sense. On account of the existing intergrades it is difficult, even impossible, to draw sharp lines between these varie- ties, and the dividing lines selected by myself (according to per- centage of diameter) may seem, and are, arbitrary: I have en- deavored, in this respect, to preserve the older names as far as possible. Sometimes it was possible to distinguish only two forms, a swollen one and a flat one; in other cases, three have been ad- mitted (an intermediate one being added to the two extremes). This latter course was made necessary in those instances, where the oldest name was given to an intergrading form. 272 ORTMANN—CORRELATION OF SHAPE AND Group OF FUSCONAIA SUBROTUNDA IN THE UPPER OHIO SYSTEM. The following is the taxonomy of this group here accepted. 1. FUSCONAIA SUBROTUNDA (Lea).—Dia. 50 per cent. of length and over. Unio subrotundus Lea. ’31 (Ohio)—Quadrula subrotunda Simp- son, ’14, p. 892.° Dia. (Lea): 60 per cent. Simpson gives: 44, 49, 51 per cent., but only the last one would belong here according to the definition accepted above. Unoi politus Say, ’34—Synonym to subrotunda according to Simpson. Dia. (Conrad; 37): 53 per cent. Fusconaia leucogona (pars) Ortmann, Nautil. 27, 713, p. 89 (Elk River, W. Va.). 2. FUSCONAIA SUBROTUNDA KIRTLANDIANA (Lea).—Dia. less than 50 per cent. Unio kirtlandianus Lea, 34 (Mahoning R., Ohio) —Quadrula kirt- landiana Simpson, 714, p. 891. Dia. (Lea) : 44 per cent.; (Simpson) : 33, 38 per cent. Fusconaia leucogona (pars) Ortmann, 1. ¢. Measurements taken from My Material. Loc. No. Max. Min. Av. Ohio River. POTESIMOTIMY cia.c ee clscraseicss see ete 5 67 51 50 subrot. IRortland parecer a meciore (les.) (les.) Soe gare wea eae oe Ot Se Sa ! SBS sBssBsesssss == (95). les. (b.-p.) b.-p. (les.) STATION IN FRESH-WATER MUSSELS. 281 These tables do not need any further comment: barring certain local irregularities, which can be accounted for by scarcity of ma- terial, the law is clearly shown, that in the upstream direction the diameter decreases, and that it decreases very gradually. The con- ditions in the Holston, and chiefly in the Clinch and Powell are, indeed, classical. It should be added that I have seen a single individual from Little Tennessee River, Coytee, Loudon Co., Tenn., which has the diameter of 58 per cent. (= pilaris). Little Tennessee goes into the Tennessee below Knoxville, and thus this agrees well with the figure for specimens from Knoxville (57 per cent.). I further collected a large number of specimens in French Broad River, at Boyd Creek, Sevier Co., Tenn. This river unites with the Holston to form the Tennessee just above Knoxville. I have measured 16 specimens; the max. is 64 per cent.; the min. 47 per cent.; the av. 56 per cent. Also these figures are exactly what we should expect. There is no question that all these shells are one and the same species, for which the name filaris, as the oldest, should be used, and which changes in obesity from the large rivers towards the headwaters. Since Lea distinguished, at the same time, two forms which differ chiefly in being more or less swollen, the two names given by him (pilaris and lesueuriana) should be preserved in a varietal sense, and a third variety of great compression should be added, described much later by Wright as bursa-pastoris. As pilaris corresponds to subrotunda, bursa-pastoris corresponds to kirtlandiana of the upper Ohio drainage. In fact, it is so close to it that Pilsbry and Rhoads’? have identified specimens from Wa- tauga River as kirtlandiana. I have serious doubts that the two forms can be kept apart, when the locality is unknown. In my large material I can see only one difference, that is, that bursa- pastoris is a smaller shell than kirtlandiana, and reaches a greater degree of compression. This opens a very pertinent question as to nomenclature, since also pilaris (and Jesuewriana) apparently repre- sent only a small race of subrotunda. I shall not go any further 12 Pr, Ac. Nat. Sct., Philad., 48, 1896, p. 502. 282 ORTMANN—CORRELATION OF SHAPE AND into this problem, which may necessitate some radical changes in our Naiad-nomenclature. This much is certain, that F. pilaris, the representative form in the upper Tennessee of F. subrotunda from the Ohio, behaves exactly as F. subrotunda, but the headwaters-forms of the two, in the upper Tennessee and the upper Ohio drainages, although very similar, have no direct genetic connection, and undoubtedly have developed independently of each other. Group oF FUSCONAIA FLAVA IN THE UPPER OHIO-DRAINAGE. I classify the forms belonging here as follows. I. FUSCONAIA FLAVA (Rafinesque).—Dia. less than 55 per cent. Obliquaria flava Rafinesque, ’20 (trib. of Kentucky, Salt, Green R.). —Quadrula flava Vanatta, Pr. Ac. Philad., ’15, p. 557. Dias (Conrad, 37): 43" per cent.; (Vanatta) 30 per cent: Unio rubiginosus Lea, ’29 (Ohio).—Quadrula rubiginosa Simpson, BI Woz: Dia. (luea’)/: 44 per cent.; (Simpson): 41, 34,53 per cent. 2. FUSCONAIA FLAVA TRIGONA’ (Lea).—Dia. 55 per cent. and over. Unio trigonus Lea, ’31 (Ohio R., Cincinnati and Louisville) — Synonym to Quadrula undata (Barnes), according to Simpson, ‘TA, p. Si. Dias (eas O2)peracent: Probably also U. undatus Barnes, ’23, belongs here. This form has been discussed by Walker (Nautil., 24, ’10, p. 24), who says that it is the same as trigonus. He gives the diameter of 68 per cent. Simpson (1. c.), who follows him, gives it as 60, 61, and 76 per cent. Walker and Simpson probably are right in regarding trigonus and undatus as synonyms, as far as it concerns obesity. However, I think that they differ in the development of the beaks, undatus having higher beaks, so that the outline is more triangular, while it is subtrapezoidal in trigonus. Of typical undatus I have no good material, and this form is not found in the upper Ohio. What I have from this region (vicinity of Pittsburgh and upwards) repre- STATION IN FRESH-WATER MUSSELS. 283 sents, in the more obese form, distinctly the shell called by Lea trigonus, and thus I shall use this name, and restrict myself to the two forms flava and trigona, and their interrelation, leaving the decision as to the standing of wndata to further investigation. Ac- cording to Utterback,’® the latter also passes into trigona, Loc. No. Max. Min. Av. Ohio River. Coraopolistesmce erase Se oats 14 63 49 54 flav. (trig.) Monongahela River. IEGI Zl CLIP average seamete sey oro lseeusiae os /otoke I 49 49 49 flav. (ClagWel cont eee eae ore eee 8 66 50 56 trig. ( flav.) IMIS ORO Misi aaceucteyete’ Occasional Papers of the Museum of Zoology 43 Obliquaria reflexa Raf., Simpson, ‘00°, p. 610; 714, p. 330. Obl. reflexa Raf., Vanatta, "15, p. 554 (“type” examined). The original description is recognizable; mentioned are the thick, convex, rounded shell, truncated posteriorly, sinuated on the post-basal margin, the rugosities of the posterior slope, and the “knobs” of the medial elevation of the shell, all char- acters of this species. O. reflexa has been designated as the type of Obliquaria Raf. by Simpson (’00). CYPROGENIA IRRORATA (Lea), 1828 Type locality: Ohio. Obovaria stegaria Rafinesque, ’20, p. 312, pl. 82, f. 4-5. Unio irroratus Lea, ’28 (not ’30 as given by Simpson), p. 260, pl. Ber heh. U. stegarius Raf.=U. irroratus Lea, Conrad, ’34, p. 71; Ferussac, 25 epee2o. .Conrad.38, p- 83, pl 40, 4 fr: Cyprogenia irrorata (Lea), Simpson, ’14, p. 320. Cypr. stegaria (Raf.) =C. trrorata (Lea), Vanatta, *15, p. 554 (“type” examined) ; Ortmann, 19, p. 218. Cypr. irrorata (Lea) = Ob. stegaria Raf., Walker, ’17°, p. 46; °18¢, p. 179 (“if identifiable”). Cypr. stegaria (Raf.), Ortmann, 718, p. 565. Rafinesque’s original figure of stegaria is absolutely insuf- ficient to recognize the species as the same as irroratus on account of the complete absence of tubercles, and, moreover, these tubercles are not mentioned in the description, except in variety (tuberculata), which is said to have a few remote tubercles; but this also does not exactly fit wrroratus, which generally has a great number of crowded tubercles. Since also nothing is said about the very characteristic color-pattern of the epidermis; except that it is brown (which it is generally not), it is impossible to identify Rafinesque’s species, and thus Lea’s name (irrorata) stands, notwithstanding the subsequent 44 University of Michigan ‘determination of the so-called Rafinesque-Poulson “type” of stegaria. “The Rafinesque-Poulson shell is stated to be of the var. tuberculata Raf. It is an ivrorata of the usual size and green- ishecolon | (al wens) Genus OpovariA Rafinesque, 1819 Type: Unio retusa Lam. Rotundaria Rafinesque, ’20, p. 308 (no type named) ; Herrmannsen, "47, p. 407 (type, Obliquaria subrotunda Raf.); Simpson, ’o0, p. 794, as a subgenus of Quadrula (type, Obliquaria tuberculata Raf.) ; Ort- mann, ’I2, p. 257, as a genus (type, the same). Obovaria Rafinesque, ’19, p. 426; ’20, p. 310 (no type named); Her- mannsen, °47, p. 132 (type, Obovaria obovalis Raf., not recognizable) ; Agassiz, 52, p. 46 (type, U. retusa Lam., congeneric with Obliq. sub- rotunda Rat.); Simpson, ’00, p. 498 (type, U. retusa Lam.). “Rafinesque proposed Obovaria in 1819 with a diagnosis but no type or recognizable species, since none of those mentioned as belonging to the genus had then been described. The diag- nosis clearly indicates a certain assemblage of Uniones which had not before been segregated, viz., those with a rounded shell with the axis (7. e., the vertical from the beaks) nearly median; such as the species subsequently included by Simpson in Obovaria, Theliderma, and Rotundaria. For some part of this assemblage the genus defined in 1819 is valid. “In 1820 Rafinesque further limited the genus, describing six species, one of which, U. obovalis, was designated type by Herrmannsen in 1847; this species has never been identified, and is believed to be unrecognizable. In 1852 another of the original species, Ob. torsa, was selected as type by Agassiz. This type has been accepted by Simpson, 1900, and by subse- quent authors, whose action is here endorsed. “As the type of Rotundaria is congeneric with O. torsa, that name becomes a synonym of Obovaria, being one year later in’ date? i(Hs A. “P2) Occasional Papers of the Museum of Zoology 45. OBOVARIA RETUSA (Lamarck), 1819 Type locality: (Incorrectly given as Nova Scotia). Unio retusa Lamarck, ’19, p. 72. Obovaria torsa Rafinesque, ’20, p. 311, pl. 82, f. 1-3. U. retusa Lam.= U. torsus Raf., Lea, ’34, p. 88 (type examined) ; Herussac;’ 35, p: 205) Conrad, 36; p. 10; pl. 8: Obovaria retusa (Lam.) = U. torsa Raf., Agassiz, ’52, p. 46. Obovaria retusa (Lam.), Simpson, ’14, p. 290. O. retusa, Clam:)' =O. torsa Raf. Vanatta; 715, “p. 552. (type” examined). Lamarck’s original description is, in spite of the incorrect locality given (Nova Scotia), perfectly recognizable. The rounded, swollen shell, with incurved beaks, and the violaceous nacre is unmistakable. Rafinesque’s torsa is also undoubtedly this, and his figures give fairly well the general character of the species. OBOVARIA SUBROTUNDA (Rafinesque), 1820 Type locality: Ohio River. Obliquaria subrotunda Rafinesque,’’20, p. 308, pl. 81, f. 21-23. Obovaria striata Rafinesque, ’20, p. 3IT. Unio circulus Lea, ’29, p. 433, pl. 9, f. 14. U. subrotundus Raf.=U. pusilla Raf.=U. striata Raf. =U. cir- culus Lea, Conrad, 734, p. 71. U, subrotundus Raf.=U. circulus Lea, Ferussac, ’35, p. 28. Obovaria circulus (Lea), Simpson, 14, p. 201. Ob. subrotunda (Raf.) =Ob. striata Raf.=Ob. circulus (Lea),. Vanatta, “15, p. 552 (“‘type” examined). Obovaria subrotunda (Raf.), Ortmann, °18, p. 567. Obl. circulus (Lea) = OdI. subrotunda Raf., Walker, ’18¢, p. 180 (“if identifiable”). Rafinesque’s description and figures indicate a rather thick, nearly round, swollen shell, with beaks almost central (“equi- laterale’), brownish-yellow epidermis, and _ whitish-purple nacre. A shell from the Ohio with these characters will be easily recognized as the circulus Lea. Thus subrotunda is. 46 University of Michigan valid. The other names given by Rafinesque (pusilla and stri- ata) do not need to be considered, since Conrad made already a selection among the available names, choosing subrotundus. OBOVARIA LENS (Lea), 1831 Type locality: Ohio and Tennessee. Unio levigata Rafinesque, ’20, p. 296, pl. 80, f. 11-13. Unio lens Lea, ’31, p. 80, pl. 8, f. Io. U. levigatus Raf. =U. lens Lea, Conrad, ’34, p. 70. Obovaria lens (Lea), Simpson, 714, p. 293. Obovaria levigata (Raf.) =O. lens (lea), Vanatta, ’15, p. 552 (“types” examined). Obovaria subrotunda levigata (Raf.), Ortmann, ’18, p. 568. Ob. lens (Lea) =U. levigata Raf., Walker, ’18¢, p. 180 (“if identi- fiable”). The description and figure of levigata indicate a strongly transversely-elliptical shell, which does not at all fit lens of Lea, which generally is rather rounded or only very slightly transverse. Thus, without knowledge of what the so-called Poulson-Rafinesque “type” is, nobody would ever suspect that Rafinesque’s shell is this. Also the character “swollen” (bombée) does not apply to lens, and levigata must be regarded as not identifiable. OBOVARIA OLIVARIA (Rafinesque), 1820 Type locality: Kentucky River. Amblema olivaria Rafinesque, °20, p. 314. Unio ellipsis Lea, ’28, p. 268, pl. 4, f. 4. U. olivarius Raf. = U. ellipsis Lea, Conrad, ’34, p. 70; Ferussac, ‘35, p. 28-34. Obovaria ellipsis (lea), Simpson, ’14, p. 299. Ob. olivaria (Raf.) =O. ellipsis (Lea), Vanatta, "15, p. 553 (“type” examined). Ob. ellipsis (Lea) = A. olivaria Raf., Walker, ’18¢, p. 180 (“if iden- tifiable’’). The specific description of olivaria may be applied to ellipsis Lea: Shell thick, little convex, oval, elliptic, beaks hardly Occasional Papers of the Museum of Zoology 47 prominent, nearly superior; epidermis striate, olivaceous ; nacre white, iridescent, length 2-3 inch. Yet it does not posi- tively indicate this species. However, in connection with the generic characters given for Amblema, the species is well characterized by the terms “dent bilobee ridee, laterale au sommet,”” which could not be used for any other species. Thus olivaria stands. Rafinesque (’20, p. 288) states in substance that he did not repeat the generic characters in his specific descriptions, as it would make them long and prolix. But when necessary, as in this case, they should be read into the description of the species. Genus ACTINONAIAS Fischer and Crosse, 1893 Actinonaias Fischer and Crosse, ’93, p. 556 (no type named, con- taining Mexican species inc. sapotalensis Lea). Actinonaias Fischer and Crosse, ’93, p. 550 (no type named, con- taining Mexican species inc. aztecorum allied to plicatulus Charp.) Nephronaias Simpson, ‘00%, p. 591 (type, plicatulus Charp.). Nephronaias Ortmann, ‘12, p. 324 (containing 3 species, incl. sapo- talensis). Nephronaias Frierson, 17, p. 47 (type, plicatulus Charp.). Actinonaias Frierson, ’17, p. 48 (type, sapotalensis Lea); Walker, "18, p. 75. The final settlement of the generic names Nephronaias and Actinonaias depends on the knowledge of the anatomy of the Mexican type of Nephronaias (plicatulus Charp.). The anat- omy of the type of Actinonatas (sapotalensis) is known (Ort- mann), and since certain North American species agree with this, this name should be used for them. The designation of the type of Actinonaias (sapotalensis) is from Frierson. ACTINONAIAS CARINATA (Barnes), 1823 Type locality: Fox River (Wisconsin). Unio crassus Say, ’17, pl. 1, f. 8 (not U. crassus Retzius, 1778). Unio ligamentina Lamarck, ‘19, p. 72. Unio crassus Say, Rafinesque, ’20, p. 293. 48 University of Michigan Unio fasciata Rafinesque, ’20, p. 204. Unio ellipticus Barnes, ’23, p. 259, pl. 13, f. 19 (not U. elliptica Rat. |°20): Unio carinatus Barnes, ’23, p. 259, pl. 11, f. Io. U. ligamentina Lam. = U. crassus Say, Lea, ’34, p. 88 (type exam- ined). U. fasciatus Raf.=U. carinatus Barn., Conrad, 734, p. 69; 735, p. Sy lk, at U. crassus Say = U. carinatus Barn., Ferussac, ’35, p. 27, 33. Lampsilis ligamentina (Lam.), Simpson, ’14, p. 79. U. crassa (Say), .Raf.=U. fasciata Raf.=—U. pallens Raf., ’°31= Lamp. ligamentina (Lam.), Vanatta, *15, p. 551 (“types” examined). U. fasciata Raf. = U. siliquoidea Bar., ’23 = U. luteolus auct., Frier- Son; “10; (1p. 130: Actinonaias ligamentina (Lam.), Ortmann, ’19, p. 232. U. crassus and U. ellipticus are nomina preoccupata. U. ligamentina is absolutely unrecognizable (the description could not possibly be poorer), and so is U. fasciata, which might be this or fasciolaris Raf. and even U. siliquoidea Barn., accord- ing to Frierson (see under L. siliquoidea), ‘The description applies to both and the so-called Rafinesque-Poulson “type”’ 1s the so-called ligamentina. It might also be any other species with elliptical outlines and with rays (iris group). The first valid name (recognizable and not preoccupied) under which the present species was published is carinatus Bar., and a good figure of this has also been given. The deter- mination of the type of U. ligamentina Lam. by Lea was sub- sequent to this, and thus has no effect. ACTINONAIAS PECTOROSA (Conrad), 1834 Type locality: Elk River, near its junction with Tennessee,. Northern Alabama. Unio vittatis Rafinesque, *31, p. 2. Unio pectorosus Conrad, ’34 (May), p. 37, pl. 6, f. 1. Unio perdix Lea, ’34 (August or September) (not ’27 as given by Simpson), -—p./ 72, “pl. 11, Hi. 31: U. pectorosus Con., ’34 (May),=U. perdix Lea, ’34 (September),. Conrad 30; pes) ple ll, tet Occasional Papers of the Museum of Zoology 49 Unio biangularis Lea, ’40, p. 288. Unio biangulatus Lea, ’42, p. 197, pl. 9, f. 8. Lampsilis biangularis (Lea), Simpson, “14, p. 59. Lampsilis perdix (lea), Simpson, ’14, p. 88. Nephronaias pectorosa (Con.) =U. perdix Lea=U. biangularis Lea = U. biangulatus lea = (possibly) U. vittatis Raf., Ortmann, ’18, p. 560. U. vittatis Raf. would have priority, if identifiable. Its shell is oval and has been compared by Rafinesque with L. fasciola Raf., but is said to be larger, rounder, with straight rays. This does not fit pectorosa, which surely is not “rounder” than fasciola, and there are additional characters in the original description which cannot be reconciled with pectorosa, chiefly “2 or 3 oblique ribs.” U. vittatis is not identifiable. It has been questionably referred to inflatus and siliquoideus Bar. by Conrad (34, p. 69). The identity of Conrad’s and Lea’s species is evident to any- one familiar with this form, and the preference of perdix by Simpson is due only to the incorrect date given for it. Already Conrad (’36) has claimed his species and has quoted the cor- rect dates. . Genus TRUNCILLA Rafinesque, 1819 Type: Truncilla truncata Raf. Truncilla Rafinesque, ’19, p. 427; "20, p. 300 (no type named) ; Herr- mansen, "40, p. 627 (type, Truncilla truncata Raf.). Amygdalonatas Crosse and Fischer, ’93, p. 557 (type, U. cognatus Lea) ; Simpson, ’00, p. 604; Ortmann, 712, p. 327. Since Tr. truncata Raf. (= elegans Lea) is supposed to be congeneric with U. cognatus Lea, and since Herrmannsen has designated truncata as the type of Truncilla, this name will have to be used for the present genus superseding that of Amygdalonaias. This will necessitate a change in the name of the genus which has been usually called Truncilla. See under Dysnomia. 50 University of Michigan TRUNCILLA TRUNCATA Rafinesque, 1820 Type locality: Ohio River. Truncilla truncata Rafinesque, ’20, p. 301. Unio elegans Wea, 31; p> $3, plo. £13: U. truncatus Raf.=U. donaciformis Lea, ’283=U. elegans ea, Conrad 344 ps 72: U. truncatus Raf.— U. elegans Lea, Ferussac, ’35, p. 27. Plagiola elegans -(1,ea), Simpson, ’I4, p. 307. Pl. elegans (Lea) =Tr. truncata Raf.=U. metaplata Raf., ’31; Vanatta, “I5, p. 553 (“types” examined). Pl. elgeans (Lea) =Tr. truncata Raf., Walker, ’16¢, p. 45; ’18¢, p. 179 (“if identifiable’). Amygdalonaias truncata (Raf.), Utterback, ’16, p. 148; Ortmann, Site, oy GyAOy This shell is small, has a “semi-triangular,’” somewhat quad- rate shape, the posterior end is truncate, and the margin and growth lines (vides) are curved (concave) behind. This is a satisfactory description of the species when compared with the only other “truncate” form of the Ohio (triquetra). U. donaciformis Lea, said to be also identical (Conrad), is closely allied, but the description of the shape does not fit it. The name 7runcilla truncata is not preoccupied by Unio truncata Spengler (1793), and thus truncata stands. TRUNCILLA DONACIFORMIS (Lea), 1828 Type locality: Ohio. Unio nervosa Rafinesque, °20, p. 296, pl. 80, f. 8-10. Unio donaciformis Lea, ’28, p. 267, pl. 4, f. 3. Umno zigzag Wea, 720, p. 440, pl. 12, f. 19. U. nervosa Raf.=U. donaciformis Lea, Say, 734. U. nervosa Raf.=U. sigsag Lea, Conrad, ’34, p. 70. U. nervosa Raf. =U. sigzag Lea, Ferussac, ’35, p. 27. Plagiola donaciformis (ea), Simpson, 714, p. 308. The original description of the rays of nervosa, “Nervures flexweuses, concentriques, vermiculaires,” might suggest U. donaciformis, yet it surely is not quite exact, and the figure, Occasional Papers of the Museum of Zoology 51 if correct in this respect, does not support this view. Further, the description says that the shell is larger behind (also shown in figure), and that the borders are undulated (see also figure), and these are characters which cannot be found in donaciformis by any means. Thus nervosa is not identifiable. Genus PLAcioLA Rafinesque, 1819 Iyvpe: Unio securis Lea == Obliquaria lineolata Raf. Plagiola Rafinesque, ’19, p. 426; '20, p. 302 (no type named) ; Herr- mannsen, "47, p. 279 (type, Obliquaria interrupta Raf.) ; Agassiz, ’52, p. 42 (first species, Obliquaria lineolata Raf. = U. securis Lea) ; Simp- son, ‘00, p. 603 (type, U. securis Lea). Herrmannsen designated an unrecognizable species Obli- quaria interrupta Raf. (see under Dysnomia brevidens) as the type of Plagiola, and consequently Plagiola should be used in the sense of Agassiz and Simpson. PLAGIOLA LINEOLATA Rafinesque, 1820 Type locality: Falls of the Ohio, at Louisville, Ky. Obliquaria (Plagiola) depressa Rafinesque, '20, p. 302, pl. 81, f. 5-7. Obliquaria (Plagiola) lineolata Rafinesque, ’20, p. 303. Obliquaria (Ellipsaria) ellipsaria Rafinesque, ’20, p. 303. Unio securis Lea, ’20, p. 437, pl. 11, f. 17. U. lineolatus Raf.=U. depressus Raf.=U. ellipsaria Raf.=U. securis Lea, Conrad, ’34, p. 70. U. lineolatus Raf. =U. depressus Raf.=securis Lea, Ferussac, ’35, p. 28. U. securis Lea=U. depressus Raf., Lea, ’38, p. 124. Plagiola lineolata Raf.=U. securis Lea, Agassiz, ’52, p. 48. U. lineolatus (Raf.), Call, ’00, p. 4609. Plagiola securis (Lea) = Obl. depressa Raf., Simpson, ’o00*¢, p. 603; "14, Pp. 305. Pl. lineolata (Raf.) = Obl. depressa Raf.= Obl. ellipsaria Rat. = Pl, securis (Lea), Vanatta, 715, p. 553 (“‘types” examined). Pl. securis (Lea) = Obl. depressa Raf. = Obl. lineolata Raf., Wal- ker, °17°, p. 45; 718¢, p. 179 (“if identifiable”). PI, securis (Lea) = Pl. lineolata (Raf.), Utterback, ’16, p. 150, foot- note. Plagiola lineolata (Raf.), Ortmann, 718, p. 571. 52 University of Michigan In the description of both depressa and lineolata Raf. the chief character of this species, great compression and posterior narrow and flat truncation, are mentioned, and thus there is no doubt about the identification. Also the peculiar color pat- tern is indicated in both. O. ellipsaria Raf. is also identifiable as this species. Conrad first pointed out the identity of Rafinesque’s species and selected the name /ineolata, and thus it has to stand. Genus LEPTODEA Rafinesque, 1820 Type: Umo fragilis Raf. Leptodea Rafinesque, ’20, p. 295 (as subgenus of Unio), no type named; Herrmannsen, °47, p. 584 (type, U. fragilis Raf.), Lasmonos Rafinesque, ’31, p. 5 (type, fragilis Raf., ’31, not fragilis, 20) ; Utterback, ’16, p. 151 (type, fragilis Raf., ’20; see below). Paraptera Ortmann, ‘II, p. 368 (type, gracilis Bar., ’23 = fragilis Raf., 20); Walker, ’18¢, p. 72. Leptodea, Frierson, ’14", p. 6 (type, leptodon Raf., ’20). Since U. fragilis Raf. has been made the type of Leptodea by Herrmannsen, Paraptera Ort. becomes a synonym having the same type, and the subsequent designation of U. leptodon Raf. as the type by Frierson becomes invalid. Utterback in giving as the type of Lasmonos the “Lasmonos fragilis Raf., ’20,’ confused Unio (Leptodea) fragilis Raf., ‘20, and Lasmonos fragilis Raf., ’°31. The latter is not identi- fiable (see under Leptodea leptodon). LEPTODEA LEPTODON (Rafinesque), 1820 Type locality: Lower Ohio River. Unio (Leptodea) leptodon Rafinesque, ’20, p. 295, pl. 80, f. 5-7. Anodon purpurascens Swainson, ’23, pl. 160. Unio velum Say, ’20, p. 293. Symphynota tenuissima Lea, ’29, p. 453, pl. 11, f. 21. Lasmonos fragilis Rafinesque, ’3I, p. 5. U. leptodon Raf.= Symph, tenuisstma Lea = An. purpurascens Sw., Conrad, ’34, p. 70. Occasional Papers of the Museum of Zoology 53 Symph. leptodon Raf. =S. tenuissima Lea= An. purpurascens Sw. =U. planus Bar., ’23, Ferussac, 35, p. 25; Conrad, ’36, p. 58, pl. 33. Lampsilis leptodon (Raf.), Simpson, ’14, p. 188. Leptodea leptodon Raf., Frierson, ’14*, p. 6. Lasmonos leptodon (Raf.), Utterback, 716, p. 156. Paraptara leptodon (Raf.), Ortmann, 18, p. 571. There is no doubt about the specific name Jeptodon Raf. and all authors have accepted it. Description and figures are entirely satisfactory. However, with regard to the validity of the generic name Lasmonos Raf., it is important to know what Lasmonos fra- gilis Raf., ’31, is. Rafinesque does not quote his Unio (Lep- todea) fragilis of ’21 under it, and thus we are to assume that it is different. The chief character of this shell is the rudi- mentary condition of the cardinal teeth, and this suggests that it might be leptodon. However, it fits also certain phases of L. fragilis (Raf.), ’20. The rest of the description does not clear up matters, since a sub-oval, thin shell, olivaceous out- side and purplish inside, fits both species, and the posteriorly broader shell with a small wing rather points to fragilis ’20. ‘ The words “some nodulities behind” fit neither. Thus Las- monos fragilis is not identifiable and Lasmonos cannot be used under any conditions. LEPTODEA FRAGILIS (Rafinesque), 1820 Type locality: Ohio River. Unio (Leptodea) fragilis Rafinesque, ’20, p. 295. Unio gracilis Barnes, ’23, p. 174. Lasmonos fragilis Rafinesque, ’31, p. 5. Unio fragilis Raf. =U. gracilis Bar., Conrad, ’34, p. 69; Ferussac, 220, p25) Conrad, ’36;p. 55, pl. 30; Frierson, “14°, p+, 7- Lampsilis gracilis (Bar.), Simpson, 714, p. 181. Lampsilis fragilis (Raf.) =L. gracilis (Bar.), Vanatta, ‘15, p. 552 (“type’”’ examined). Lasmonos fragilis Raf., Utterback, 716, p. 152 (quotations in part incorrect ). 54 University of Michigan Paraptera fragilis (Raf.) =U. gracilis Bar., Ortmann, 18, p. 572. Lamp. gracilis (Bar.) =U. fragilis Raf., ’20=Lasmonos fragilis Raf., °31; Walker, ‘18¢, p. 182 (“if identifiable”). The description of U. fragilis, chiefly its comparison with U. leptodon, makes it sure that this is the gracilis Bar. The shell is thin, fragile, not elongate and attenuated posteriorly (as leptodon is), but somewhat dilated behind. Also the char- acters of the pseudocardinals are mentioned. It remains doubtful whether Lasmonos fragilis is this shell or Leptodea leptodon (see under the latter). Genus CARUNCULINA Simpson, 1898 Type: Unio parvus Barnes. Toxolasma Rafinesque, °31, p. 2 (no type named). Corunculina Simpson, ’98, p. 109 (error typographicus) (only spe- cies, U. parvus Bar.). Carunculina Simpson, ‘co’, p. 563, and ’14. p. 148 (type, Unio tex- asensis Lea). Toxolasma Raf., Frierson, ’14*, p. 7 (type, lividus Raf. =glans Lea); Ortmann, 718, -p. 572: The revival of the generic name To.volasma depends upon the identity of U. lividus Raf. As will be shown under Car. mosta, lividus is not recognizable, and thus the name T0.ro- lasma should be discarded. As type of Carunculina, U. parvus must be taken, for at the first publication of this subgenus only this species was included. CARUNCULINA MasTA (Lea), 1841 Type locality: French Broad River, East Tennessee. Unio (Toxolasma) lividus Rafinesque, ’31, p. 2. Unio pullus Conrad, ’38,.p. 100, pl. 55, f. 2. Unio mastus Lea, 41, p. 82. Unio cylindrellus Lea, ’68, p. 144. Unio corvunculus Lea, ‘68, p. 144. Unio glans Pilsbry and Rhoads, ’96, p. 502 (not glans Lea, ’34). Lampsilis cylindrella (Lea), L. masta (Lea), L. corvunculus (Lea), L. pullus (Con.), Simpson, ’14, pp. 155-160. Occasional Papers of the Museum of Zoology 55 Toxolasma lividum (Raf.) = U. glans Lea, Frierson, 14", p. 7. Toxolasma lividum (Raf.) =U. pullus (Con.) (part) = U. mestus Lea = U. cylindrellus Lea, Ortmann, ’18, p. 573. U. (Tox.) lividus Raf.=Car. glans (Lea) or possibly = pullus Con., Walker, ’18¢, p. 180-181 (“if identifiable’). This is the form representing Car, glans Lea in the upper Tennessee region, but it differs from the typical glans of the interior basin at least as a variety. It has been claimed by Frierson and Ortmann that U. lividus Raf. is this form. It is, according to description, a small shell (1 inch) of elliptical shape, “swelled,” not thick, with rough, brown epidermis and livid purple nacre, from Rock Castle River, Kentucky (Upper Cumberland). This description may be applied to two species of this region, the present one and Ligumia vanuvemensis (Lea), and there is nothing in it which permits a final decision. Moreover, the dimensions given by Rafinesque fit in part (diameter 37% of length) both species, in part (height 75%) neither of them. Thus lividus is not recognizable and cannot be used, and with it the generic name 7orolasma must be rejected. Occasional specimens of vanu.remensis have quite exactly the proportions given by Rafinesque. _U. pullus Conrad resembles this form very much. But since it originally came from an entirely different region ( Wateree River, South Carolina) it cannot be identified with certainty and should be disregarded, at least for the present. U. mestus Lea surely is this upper Tennesee form of glans (from French Broad River). The figure represents a very large male ; but similar specimens have been found by Ortmann in the French Broad drainage, fully agreeing with this in all characters, including the proportional dimensions (height of male, 58-63% of length; of females, 63-08% ; diameter of male 38-43% ; of female, 39-48% ; Lea’s figures for height and 56 University of Michigan diameter are 60% and 40%, respectively). Thus the name mastus Lea is the oldest available one. The later names given by Lea and claimed by Ortmann as possible synonyms (cylindrellus and corvunculus) need not be considered here. Their validity depends upon taxonomic con- siderations, and these two forms require additional study before their standing can be settled. Genus CoNRADILLA Ortmann, 1921 Type: Unio celatus Conrad. Conradilla Ortmann, Naut., XXXIV, 1921, p. go. CONRADILLA C&LATA (Conrad), 1834 Type locality: Tennessee, Elk, and Flint rivers. Unio (Lemiox) rimosus Rafinesque, ’31, p. 3. Umo celatus Conrad, ’34, p. 338, pl. 1, f. 2; ’34, p. 20, pl. 13, f. 4. Micromya celata (Con.), Simpson, ’14, p. 34. Lemiox rimosus (Raf.) =U. celatus Con., Frierson, ’14, p. 7. Lemiox rimosus (Raf.), Ortmann, ’16, p. 39; '18, p. 574. Micromya celata (Con.), Walker, 18°, p. 4, 18°, p. 185 (new generic name should be proposed). Frierson has identified U. rimosus with U. celatus chiefly on the strength of the word “rimose”’ in the description, which is intended to describe the sculpture of the shell. Walker objects to this. Although this word may be taken as giving a good description of this feature of the shell, other points in the description do not fit. Rafinesque (731, p. 4) makes the following explicit state- ment of his theory of comparative measurements: “The com- parative proportions of the length, breadth, diameter and axis of the Unios and other bivalve shells having been misunder- stood by some, it may be needful to state that my formula is a kind of abbreviation of a longer exposition. Thus, when I say, length one-half, diameter one-third, axis one-fourth of the breadth, | meant to say and must be understood to state Occasional Papers of the Museum of Zoology 57 the following longer account: The length of the shell is one- half, the diameter is one-third, and the axis is at one-fourth of the breadth, or largest dimension of the shell. “In longitudinal shells this is reversed, the length being the longest dimension becomes the size of comparison.” Vanatta (15, p. 549) seems to have overlooked this state- ment, and consequently his explanation of the meaning of Rafinesque’s fractions is obviously incorrect. In course of the investigations made in the preparation of this paper we have found Rafinesque’s comparative measure- ments in most cases very exact and are of the opinion that they are to be relied upon as a means for identifying many of his species. The discrepancy pointed out by Walker (18, p. 5) 1s an important one and a serious objection to the approx- imation of rumosus to this species. But this much is sure, that Rafinesque describes his shell as elliptic, which c@latus is not, and that he describes it as broader behind, which again does not fit. It is also correct that the “rimose”’ character of the shell in celatus is not restricted to the posterior part, while the main part of the surface is smooth, as stated in the description of rimosus, but that it covers nearly three-fourths of the entire shell. The term “rimose” might also be applied, according to Wal- ker, to Medionidus conradicus (Lea), and the rest of the description would not speak against this. “In discussing U. rimosus Raf. Frierson claims that by ‘Broader . . . behind’ Rafinesque means what we would term longer behind; but even so, the dimensions do not fit. Rafinesque’s shell measured, L: 11%4, H.1, D. % inch. A speci- men measures, L. 1%, H. 1%, D. % inch. This is a rather compressed and low example, but more than double the diam- eter assigned by Rafinesque. It may be claimed that Rafin- 58 University of Michigan esque’s measurement of diameter was an error; in cases where a figure is present to serve as guide for such corrections they are justified. Otherwise nobody has a right to change an author’s express statements to make them conform to a theo- retical identification. Unio rimosus is not identifiable.” (H. As sles) Thus, the name rimosus for the species cannot be used, and consequently also Lemiox for the genus is not available. Since the species should be placed in a separate genus (see Ortmann ) a new name was needed, and Ortmann (1. c.) has proposed that of Conradilla. MEDIONIDUS CONRADICUS (Lea), 1834 Type locality: Unknown. Unio platcolus Rafinesque, ’31, p. 3. Unio conradicus Lea, 34, p. 63, pl. 9, f. 23. Unio conradicus Conrad, °38, p. 87, pl. 47, f. 3. Medionidus conradicus (Lea), Simpson, “14, p. 247; Walker, ‘18¢, p. 70. Medionidus plateolus (Raf.) = conradicus Lea, Ortmann, ’18, p. 575. The original description of U. plateolus Raf. is entirely insufficient to recognize this species. The chief characters are: general shape (elliptic-lanceolate and very compressed), small size (2 inches), and color (brown, inside bluish). They apply to conradicus, but also to a number of other species, chiefly of the genus Liguimia (trabalis and the iris group), or a young leptodon, or a flat, young dilatata. Besides, the shell is called attenuate and subacute behind, which does not fit conradicus very well, and the characteristic wrinkles of the posterior slope in this species and the rays, which generally are distinct, have not been mentioned. Thus plateolus is not identifiable. Occasional Papers of the Museum of Zoology 59 Genus LicuMra (Swainson), 1840 Type: Unio recta Lam. Eurynia Rafinesque, ’20, p. 297 (no type named) ; Herrmannsen, °47, p- 564 (type, U. dilatata Raf.) ; Agassiz, ’52, p. 45 (type, U. recta Raf.) ; Simpson, ’00, p. 534 (type, U. recta Lam.). Ligumia Swainson, °40, p. 378 (type, recta Lam.). The first designation of a type for Eurynia was that of U. dilatata Raf. by Herrmannsen, and thus Eurynia becomes a synonym of Elliptio (which see). This opens the way for the admission of Ligumia Swainson with the same type as Eurynia of Agassiz and Simpson. LIGUMIA REcTA (Lamarck), 1819 Type locality: Lake Erie. Unto recta Lamarck, ’19, p. 74 (Lake Erie). U. rectus Lam. = U. latissimus Raf., Conrad, ’34, p. 71; Ferussac, 35D. 27) Manatta, “Sep. 55l © type examined): Unio sageri Conrad, °36, p. 53, pl. 29, f. 1 (Detroit River). Lampsilis recta sageri (Con.), Simpson, ‘14, p. 96 (Lake Erie and Detroit River). Eurynia recta (Lam.), Ortmann, ‘18, p. 582 (Lake Erie) ; Walker, "18¢, p. 184 (Lake Erie). LIGUMIA RECTA LATISSIMA (Rafinesque), 1820 Type locality: Ohio River. Unio latissima Rafinesque, ’20, p. 297, pl. 80, f. 14-15 (Ohio River). Unio rectus Conrad, 736, p. 33, pl. 15 (interior drainage) (not U. recta Lam.). Lampsilis recta (Lam.), Simpson, ’14, p. 95 (Mississippi drainage). Eurynia recta latissima (Raf.), Ortmann, ‘18, p. 582 (interior drain- age) ; Walker, ’18¢, p. 184 (Ohio drainage). The statement of Conrad, Férussac, and Vanatta that recta Lam. is identical with lJatissima is correct only when we do not separate these two forms, but if they should be separated as varieties, recta Lam. refers to the Lake Erie form, which is evident not only from the locality given but also from the 60 University of Michigan description of the color of the epidermis: blackish-brown. The Ohio form is pure black or greenish-black. Rafinesque’s Jatissima is well described, the elongated shape, black epidermis, and large size being mentioned, and undoubt- edly refers to the Ohio form. U. sageri thus becomes a syno- nym of recta, which is the main species, and latissima is the variety. LAMPSILIS ANODONTOIDES (Lea), 1831 Type locality: teres, Wabash River; anodontoides, Missis- sipp1, Alabama, and Ohio rivers. Unio teres Rafinesque, ’20, p. 321. Unio anodontotdes Lea, ’31 (not ’34, as Simpson gives), p. 81, pl. 8, Paes Unio teres Raf. =anodontoides Lea, Conrad, ’34, p. 72; Ferussac, °35, p. 27; Conrad, ’36, p. 52, pl. 28 (Poulson’s “type” examined and figured). U. teres Rat., ‘Call. 00, pi 452. Lampsilis anodontoides (Lea), Simpson, ’14, p. 90, and L. fallaciosa (Smith) (799) Simpson, ’14, p. 92. Lampsilis anodontoides (Lea) =U. teres Raf., Utterback, ’16, p. 179, foot-note. “Whether the Unio teres Raf. was based upon specimens of U. anodontoides Lea or Lamp. fallaciosa Smith is a question upon which authorities of equally good judgment have held opposing views and where certainty does not seem attainable. Conrad’s figure, said to be from a specimen labelled by Rafin- esque, is anodontoides; but it is larger than Rafinesque’s meas- urements, therefore not the original type. It has been lost. “Rafinesque’s type measured ‘envirow L,. 75, H. 30, D. 50 mm. ‘The nearest specimen now measured is, L. 78, H. 33, D. 24 mm. In the most obese examples of either species the height still surpasses the diameter, which in Rafinesque’s shell was said to be far greater than in any specimen of either spe- cies mentioned. Occasional Papers of the. Museum of Zoology 61 “As stated above, the method of ‘correcting’ an author’s measurements to force them to agree with what species we will is essentially unscientific. It savors of medizval theolog- ical procedure. If great dependence is placed upon these pro- portions in some cases, why throw them away when they do not suit our convenience? In the case of Unio teres, its adop- tion would mean the rejection of certainties for a name resting on a description requiring arbitrary alteration, and after that, practically arbitrary selection between two closely related forms. U. teres should be discarded as not identifiable.” (H. ASCE) LAMPSILIS SILIQUOIDEA (Barnes), 1823 Type locahty: Wisconsin River. Unio luteola Lamarck, 719, p. 79. Unio fasciata Rafinesque, ’20, p. 204. Lampsilis fasciola Rafinesque, ’20, p. 299. Unio inflatus Barnes, ’23, p. 266. Unio siliquoideus Barnes, '23, p. 269, pl. 13, f. 15. U. cariosus Say, ’17= U. luteola Lam., Lea, ’29, p. 417-425; Conrad, "34, p. 68; Ferussac, 735, p. 26. U. siliquoideus Bar. = U. inflatus Bar., Lea, ’29, p. 419; Conrad, *36, pp. 23-40. U. inflatus Bar. = U. siliquoideus Bar., Conrad, ’34, p. 69. Us luteola wam. =U. siliquoideus Bar., Lea, 734, p. of (type exam- ined). U. fasciata Raf. =U. carinatus Bar., ’23, Conrad, ’34, p. 69; 735; p. aepleui. U. fasciolus Raf.=U. multiradiatus Lea, ’29, Conrad, ’36, p. 26, pl. 11, f. 2 (Poulson’s specimen examined). U. siliquoideus Bar. = U. inflatus Bar., Conrad, ’36, p. 22, pl. 9, f. I. Lampsilis luteola (Lam.), Simpson, ’14, p. 60. Lamp. luteola (Lam.) = L. fasciata Raf., Vanatta, ’15, p. 551 (“type” examined). U. fasciata Raf.=U. siliquotideus Bar.=U. luteolus Lam. (of authors), Frierson, *19, p. 139. Of the names applied to this form, the oldest is Juteola Lam. According to description and locality this has been referred to 62 . University of Michigan U. cariosus Say ’17, by Lea, Conrad, and Férussac, but Lea changed his opinion after the examination of Lamarck’s type, identifying it with siliquoideus Bar. There is no question that Lamarck’s brief description applies better to cariosa (“‘poste- rior end broader and rounded” ) and that of the localities given, the first (Susquehanna) has only cariosus, but not siliquoideus. If the type of luteola is the latter, the description is unsatis- factory. In either case, the name /wfeola cannot be used. The next name to be discussed is fasciata Raf. Conrad con- siders this to be carinatus Bar. (ligamentinus auct.), and according to Vanatta the “type” confirms this. However, Frierson thinks that it is siliquoideus. The original description of fasciata (see also under Actinonatas carinata) gives the characters: elliptic, convex, shell not thick; epidermis little rugose, olivaceous, with brown rays; nacre bluish, cardinal tooth rugose, divaricate ; lateral tooth carinate; size up to over three inches. This fits both species, and thus fasciata is not identifiable. In view of Rafinesque’s statement that his fasciata occurs in the Ohio, Muskingum, Kentucky, Salt, Green, and other rivers, and that while it is ordinarily a small species, yet he had seen it more than three inches in length, it would seem to be conclusive that it is not the same as siliquoidea Bar. A series of comparative measurements shows that of the possible species known in the Ohio system to which it might be approximated fasciola Raf. and carinata gibba Simp. are the only ones that at all correspond to those given by Rafin- esque. As between these two, so far as proportions are con- cerned, there is not much choice. In the absence of any speci- fication as to the character of the rays of fasciata, it is impos- sible to refer it with any certainty to either, although the com- parison with ochracea would seem to indicate fasciola rather Occasional Papers of the Musewm of Zoology 63 than carinata gibba. It must, therefore, be held to be uniden- tifiable. Then follows U. fasciola Raf. According to Vanatta, the Poulson type belongs to luteola auct. (= siliquoidea Bar.), but this is contradicted by Conrad, who mentions also a speci- men from the Poulson Collection, which is called fasciolus, and is identical with multiradiatus Lea. Moreover, as will be shown under Lampsilis fasciola, Rafinesque’s description does not fit the present species, but does fit multiradiata very well. Thus, also, this name cannot be used. Next are two names given by Barnes, ’23, inflatus and sili- quoideus. That these are identical and belong here has been recognized by Lea, ’29, confirmed by Conrad, and accepted by Simpson, and there is no doubt that siliqguoideus is the male, imflatus the female. Of these two names, siliqguoideus has been selected by Lea in ’29, while Conrad, in °34, selected inflatus, but changed subsequently (’36) to siliquwoideus. The latter thus is the valid name, and should be used for the species hitherto commonly called Jiuteola. LAMPSILIS VENTRICOSA (Barnes), 1823 Type locality: Wisconsin and Mississippi rivers, Prairie du Chien, Wis. Lampsilis cardium Rafinesque, 20, p. 298, pl. 80, f. 16-10. Unio ventricosus Barnes, '23, p. 267, pl. 13, f. 14. U. cardium Raf. = U. ventricosus Bar., Conrad, ’34, p. 68 Lampsilis ventricosa (Bar.), Simpson, ’14, p. 38. L. cardium Raf.=L. ventricosa (Bar.), Vanatta, ’15, p. 551 (“type”’ examined ). L. cardium Raf.=L. ovata Say, °17, Ortmann, ’18, p. 583. L. ventricosa (Bar.) =L. cardwim Raf., Walker, ’18¢, p. 184 (“if identifiable” ). Conrad and Vanatta (examination of the alleged Poulson- Rafinesque “type”) make cardimm the same as ventricosa. 64 University of Michigan From the original description this cannot be confirmed. Car- dium might be ovata, ventricosa, or even Proptera capax Green, and from the comparison of L. ovata Say with cardium given by Rafinesque we should conclude that cardium is the female of ovata (ovata differs chiefly by the less swollen shape and non-dilated posterior end). This is also supported by the figure of cardium, which shows distinctly a rather sharp pos- terior ridge. At the best, cardiuwm is not identifiable, and thus. this name cannot supersede ventricosus. LAMPSILIS FASCIOLA Rafinesque, 1820 Tvpe locality: Kentucky River. Lampsilis fasciola Rafinesque, ’20, p. 290. Unio multiradiatus Lea, ’29, p. 434, pl. 9, f. 15. U. fasciolus Raf.=U. multiradiatus Lea, Ferussac, ’35, pp. 26, 32; Conrad, ’36, p. 26, pl. 11, f. 2 (Poulson’s specimen examined). Lampsilis multiradiata (Lea), Simpson, ’14, p. 55. Lampsilis luteola (Lam.) =L. fasciola Raf., Vanatta, ’15, p. 551 (Poulson’s “type” examined). L. fasciola Raf. =U. multiradiatus Lea, Ortmann, 718, p. 584. Conrad and Vanatta have examined a specimen in the Rafin- esque-Poulson Collection, and Férussac an authentic specimen from Rafinesque ; Conrad and Férussac pronounce fasciola to be the same as multiradiata, while Vanatta says that it is lute- ola (= siliquoidea). There is evidently some mistake about the supposed “type.” However, Rafinesque’s description is unmistakable. It “indi- cates a shell of the cardium-ovata type, with unequal, flexuous rays, which fits multiradiata Lea, but not luteola Lam.” (Ort- mann), and Conrad has also pointed out as the essential char- acter the numerous “unequal, green, undulated or flexuous rays.” Thus fasciola is identifiable and valid. Occasional Papers of the Museum of Zoology 65 Genus Dysnomra Agassiz, 1852 Type: Unio foliatus Hild. = Obliquaria flexuosa Raf. Truncilla Rafinesque, ’20, p. 300 (no type named); Herrmannsen, "47, p. 627 (type, Truncilla truncata Raf.) ; Agassiz, ’52, p. 44 (no type named, first species, Tr. triqueter Raf.) ; Simpson, ’00, p. 516 (type, Tr. triqueter Raf.). Dysnomia Agassiz, '52, p. 43 (no type named, first species, Obli- quaria flexuosa Raf.) ; Simpson, ‘00, p. 521, as subgenus of Truncilla (type, U. foliatus Hild. = Obl. flexuosa Raf.). Since the type of Truncilla has been designated by Herr- mannsen as Tr. truncata Raf., this name takes the place of Amygdalonaias (which see). The next available name for this genus is Dysnonua Agassiz, the type of which has been fixed by Simpson. Subgenus TRUNCILLOPSIS n. n. Type: Truncilla triqueter Raf. Truncilla (subgenus) Simpson, ‘00, p. 517 (type, Truncilla triqueter Raf.). The removal of 7runcilla and the extension of the subgen- eric name Dysnomia to cover the whole: genus necessitates the introduction of a new subgeneric name for what was hitherto the subgenus Truncilla s. s. of Simpson. The name of Trun- cillopsis is therefore proposed. DysNoMIA (‘TRUNCILLOPSIS) TRIQUETRA (Rafinesque), 1820 Type locality: Falls of the Ohio (at Louisville, Ky.). Truncilla triqueter Rafinesque, ’20, p. 300, pl. 81, f. 1-4. Unio triangularis Barnes, ’23, p. 272, pl. 13, f. 17. Unio cuneatus Swainson, Phil. Mag., ’23, p. 112. Unio formosus Lea, °31 (not ’34, as given by Simpson), p. 111, pl, TOM te Sus U. triqueter Raf. =U. triangularis Bar.=U. formosus Lea=U. cuneatus Sw., Conrad, ’34, p. 72; Ferussac, ’35, p. 27. Truncilla triquetra Raf., Simpson, ’14, p. -Truncilla triquetra Rat. Vanatta, “15, p: oat 50 (“type” examined). cn 66 University of Michigan The generally accepted identity of Tr. triquetra Raf., which is evident from description and figure, has been confirmed by Vanatta’s examination of the so-called Rafinesque-Poulson Stype-- DysNomiIA (TRUNCILLOPSIS) BREVIDENS (Lea), 1831 Type locality: (Incorrectly given as Ohio) corrected by Lea, 34, p. 85, to Cumberland River. Obliquaria interrupta Rafinesque, ’20, p. 302. Unio brevidens Lea, ’31 (not ’34, as given by Simpson), p. 75, pl. 6, tO: U. interruptus Raf. = U. brevidens Lea, Conrad, ’34, p. 69; Ferussac, 35, p. 28; Conrad, ’38, p. 88, pl. 48. _ Truncilla brevidens (1.ea), Simpson, ’14, p. 7. Truncilla brevidens (Lea) = Obl. interrupta Raf., Vanatta, ’I5, p. 550 (“type”’ examined). Tr. brevidens (lea) = Obl. interrupta Raf., Walker, ’16¢, p. 45; ’18¢, p. 186 (“if identifiable”). Tr. interrupta (Raf.), Ortmann, ’18, p. 586. Vanatta’s view. that Obliquaria interrupta Raf. is preoccu- pied by Unio solenoides interrupta Raf. does not hold good (Walker). However, the original description of O. interrupta is not sufficient to recognize the species. It is evident that Rafinesque’s type of his interrupta, if a specimen of brevidens, was a male shell, no mention being made of the characteristic posterior inflation and truncation of the female brevidens. A normal male brevidens of exactly the same length as the type of imterrupta is 10% higher and 25% more inflated. Many specimens could be selected that are proportionately higher. The proportions given by Rafin- esque for his shell do not, therefore, agree with those of brevidens. The question of locality should also be considered. Rafin- esque says, “Found in the Kentucky and Ohio rivers.” Sa far as we have been able to ascertain, there is no record of the Occasional Papers of the Museum of Zoology 67 occurrence of brevidens in the Ohio. It is not given in any of the Ohio lists that we have seen. Brevidens is a character- istic species of the Tennessee system. Like some other Ten- nessee species, by stream transference in the head-waters, it has got into the Cumberland River, but not out into the Ohio. We know practically nothing of the fauna of the Kentucky. We have no records of this species having been found there. On the other hand, the various forms of the nebulosa group are generally distributed in the Ohio and its southern tribu- taries. It seems very strange that Rafinesque seems never to have collected any species of that group. At least none have been identified among his species. An “ovate-elliptical shell, not thick, and little swollen, with reddish-brown epidermis and interrupted rays and white nacre,” may be very well applied also to forms of Ligumua of the nebulosa-iris group. In this connection attention should also be called to the possibility that Rafinesque’s shell may have been a specimen of teniatus Con. A specimen of nearly the same size has exactly the same proportions as those given by Rafinesque for his shell. In all other respects, except perhaps color, including the character of the lateral teeth, it agrees with interruptus. While this species does not occur in the Ohio, it is found in the Cumberland and is quite likely to be found in the Kentucky as brevidens. Punctata Lea would also apply. Except that it is usually not so high proportionately as mterruptus, in other respects it agrees quite as well as teniata. It would seem to us that it is quite as possible that Rafinesque had one of these or some allied species before him as that he had brevidens. “The Rafinesque-Poulson specimen of interruptus is a female brevidens, very solid, diameter nearly one-half the length, 68 University of Michigan 55-5 x 26.5 mm.; ‘pew epats’ would alone throw it out.” (H- AGS) Subgenus SCALENILLA n. n. Type: Unio sulcatus Lea. Scalenaria Rafinesque, ’20, p. 309 (no type named) ; Herrmannsen, "48, p. 422 (type, Obliquaria scalenia Raf.) ; Agassiz, ’52, p. 43 (no type named) ; Simpson, ‘00, p. 519 (type, Unio sulcatus Lea). The type of Scalenaria has been designated by Herrmann- sen as Oblig. scalenia Raf., which is the same as Pleurobema clava (Lam.), which see. Thus this name becomes a synonym of Pleurobema. Subsequent attempts by Agassiz (first spe- cies) and Simpson to select, as type Obliq. obliquata Raf. (an. unidentifiable species) or U. sulcatus Lea (supposed to be the same as obliquata) are invalid. Since there is no avail-— able name for this subgenus, we propose that of Scalenilla. DysNOMIA (SCALENILLA) SULCATA (Lea), 1829 Type locality: Ohio, Obliquaria (Scalenaria) obliqtata Rafinesque, ’20, p. 309 (Kentucky River). Unio sulcatus Lea, “May or June,’ 1829 (not 730, as given by Simp- son); p. 430, ‘pl. °8,. t. 12. Unio ridibundus Say, October 7, 1820, p. 308. U. obliquatus Raf.= U. sulcatus Lea, Conrad, °34, p. 70. U. obliquatus Raf.—=U. sulcatus Lea = U. haysianus Lea =U, ridi- bundus Say, Ferussac, 35, pp. 28, 34. Scalenaria obliquata (Raf.) = U. sulcatus Lea (male) = U. ridibun- dus Say (female), Agassiz, °52, p. 43. Truncilla (Scalenaria) sulcata (lea), Simpson, ’14, p. 14. Trunc. obliquata (Raf.) =Tr. sulcata (Lea), Vanatta, ’15, p. 550+ (“type” examined). Tr. sulcata (Lea) = Ob. obliquata Raf., Walker, ’16°¢, p. 45, and ’18¢,. p. 186 (“if identifiable”). U. sulcatus Lea is not preoccupied by Pleurobema cuneata sulcata Raf. ((20), as Vanatta claims (see Walker). Obl. obliquata is not identifiable from the original descrip-- Occasional Papers of the Museum of Zoology 69 tion, for every word of it applies as well to Pleurobema pyra- midatum (lea): shell thick, swollen, ovate-triangular, the three sides curved, a light oblique and longitudinal depression ; epidermis nearly smooth, black; nacre rose-purplish; size 2-3 inches. In fact, the black epidermis and the size fit U. pyra- midatus better than U. sulcatus. Thus the name obliquata cannot be revived. DysNoMia (PILEA) TURGIDULA (Lea), 1858 Type locality: Cumberland River and Florence, Ala. Unio turgidulus Lea, ’58, p. 40 (male). Unio deviatus Reeve, ’64, pl. 15, f. 61 (female). Truncilla deviata (Rve.) (male and female) Walker, ’10°, pp. 78, 81. Tr. florentina (1,ea) (in part) and Tr. deviata (Reeve), Simpson. "14, pp. 30-31. Tr. curtisi Frierson and Utterback, Utterback, ’16, p. 190, pl. 6, f. 14, DleezSnata loo: Tr. turgidula (Lea) = U. deviatus Rve., Ortmann, ’18, p. 590. Tr. curtisi Fr. and Utt.= Tr. deviata (Rve.), Walker, ’18¢, p. 185. Walker (10) was the first to indicate that turgidulus Lea is the male of the female deviatus, by arranging his key for the males of Truncilla in such a way that the characters seen in turgidulus lead to the male of deviatus, but he did not expressly stated this in the text, using only the name deviata, while turgidulus has the priority. The identity of Tr. curtist from the Ozarks has been recog- nized by Walker (’18) and is evident from Utterback’s description and figures. The examination of authentic speci- mens distributed by Utterback has settled this beyond any doubt. DysNOMIA (PILEA) TORULOSA (Rafinesque), 1820 Type locality: Ohio and Kentucky River. Amblema torulosa Rafinesque, ’20, p. 314, pl. 82, f. 11-12. Amblema gibbosa Rafinesque, ’20, p. 315. 70 Unizersity of Michigan Unio perplexus Vea, ’31, p. 112, pl. 27, £. 42. U. torulosus Raf.= U. gibbosus Raf.=U. perplexus Lea, Conrad, 34, P 72. U. gibbosus Raf. =U. perplexus Lea, Ferussac, ’35, p. 27. U. gibbosus Raf.=U. torulosus Raf.—=U. perplexus Wea, Conrad, 730,;-P 550, Dl 27, teat: Truncilla perplexa (Lea), Simpson, 714, p. 24. Tr. torulosa (Raf.) = Ambl. gibbosa Raf.—=Tr. perplexa (Lea), Vanatta, “15, p. 550 (“type” examined). Tr. torulosa (Raf.), Ortmann, ’18, p. 580. Tr. perplera (Lea) = A. torulosa Raf. = A. gibbosa Raf., Walker, *18¢, p. 186 (“if identifiable”). “Lea’s Unio perplexus is preoccupied by Say for a variety of U. ridibundus not noticed by Simpson (Am. Con., Pt. I, Binney s. Reprint ps 155). (Cele:As.E:) Rafinesque’s crude figure of A. torulosa is sufficient to rec- ognize in it the female of this species, since it distinctly shows the posterior expansion of the female shell and the character- istic nodes. Also the description confirms the identity. 4. gibbosa undoubtedly is the male belonging here, and the two large, nodulous ribs and the oblique furrow between them are unmistakable. This identification with perplerus Lea is sup- ported by Vanatta’s examination of the so-called Rafinesque- Poulson “type.” Conrad’s first selection (°34) of the name torulosa in pref- erence to gibbosa must stand, although he subsequently (’36) reversed this. DysNoMIA (DyYSNOMIA) FLEXUOSA (Rafinesque), 1820 Type locality: Kentucky, Salt, and Green rivers. Obliquaria fleruosa Rafinesque, '20, p. 309. Unio foliatus Hildreth, ’28, p. 284, p. 16. Unio (Epioblasma) biloba Rafinesque, *31, p. 2. U. gtbbosus Raf. probably = Epioblasma biloba Raf., Ferussac, ’35, Pp. 27-34 (authentic specimen of biloba examined). U. flexuosus Raf. = foliatus Hild., Conrad, 35, p. 8, pl. 4, f. 2 (‘‘type” examined). Occasional Papers of the Museum of Zoology 71 Dysnomia flexuosa (Raf.) =U. foliatus Hild., Agassiz, ’52, p. 43. Truncilla (Dysnomia) foliata (Hild.), Simpson, ’14, p. 18. Epioblasma biloba Raf.=U. foliatus Hild., Frierson, ’14*, p. 7. Trun, flexuosa (Raf.) =Tr. foliata (Hild.), Vanatta, ’15, p. 557 (“type” examined). Tr. foliata (Hild.) = Obl. flexuosa Raf.= Ep. biloba Raf., Walker, "18¢, p. 186 (“if identifiable”). Conrad and Vanatta have found that the so-called Rafin- esque-Poulson “type” of fexuosa is the same as foliatus Hild. The original description of feruosa mentions two gentle ele- vations of the shell, and between them a wide and flat depres- sion. This is a very prominent character of the species and cannot be applied to any other shell. Also the description of the margins of the shell as “‘flexuous” is significant. Thus the specific name f#exuosa stands. As to biloba, its recognition is important with regard to the validity of the subgenus Epioblasma (as against Dysnomia). Férussac is not sure about the identity of his authentic speci- men of biloba. The phrase “‘belly bilobed” has suggested the idea that it is the female of f#eruosa, but this is not bilobed below, but bisinuata, with only one large lobe. Also the description of the shell as elliptical does not fit, for it is tri- angular. Further, the beaks are not prominent, as described, and nothing is said about the very striking feature, the two ridges of the shell, and the deep and wide radial furrow. Thus the description of biloba is not recognizable, and with this name also that of the subgenus, Epioblasma, goes into the discard. 72 1817-1819 1819 1819 1828 1829 1829 1831 1830-1834 1831° 1831 1832 1832 University of Michigan BIBLIOGRAPHY Say, J., Conchology in Nicholson’s Encyclopedia, eds. 1, 2, 3, 1817-1819. LAMARCK, J. B. be, Historie naturelle des Animaux sans vertebres, VI, 1819. RAFINESQUE, C. S., Prodrome de 70 nouveaux Genres, etc. (Journ. de Phys., Brux., pp. 423-428). RAFINESQUE, C. S., Monographie des Coquilles Bivalves et Fluviatiles de la riviere Ohio (Ann. Sci. Phys. Bruxelles, V, 1820, pp. 287-322). SwaAInson, W., Zoological Illustrations, II, 1821-2, Barnes, J. W., On the genera Unio and Alasmodonta: with introductory remarks (Amer. Journ. Sci., VI, 1823, pp. 107-127, 258-280). GREENE, J., Some remarks on the Unios of the United States, with a description of a new species (Contrib. Mac- lurian Lyc., I, 1827, pp. 41-47). VALENCIENNES, A., Recueil d’Obs. de Zool. et d’Anat. Comp., WolA1ts HinpretH, S. P., Observations on and descriptions of the shells found in the Muskingum River (Amer. Journ. Sci., XIV, 1828, pp. 276-291). Lea, IL, Descriptions of six new species of the genus Unio, etc. (Trans. Amer. Philos: Soc., III, 1828, pp. 259-273). Lea, I., Description of a new genus of the family Naiades, including eight species, four of which are new; also the description of eleven new species of the genus Unio, etc. (Trans. Amer. Philos. Soc., III, 1829, pp. 403-457). Say, J., Description of new terrestrial and fluviatile shells of North America (New Harmony Disseminator, 1829). Lea, I., Observations on the Naiades, and Descriptions of new species of that and other families (Trans. Amer. Philos. Soc., IV, 1831, pp. 63-121). Say, T., American Conchology. Say, T., Description of several new shells (Transylvania Journal, IV, 1831). RAFINESQUE, C. S., Continuation of a Monograph of the Bivalve shells of the River Ohio, 1831, pp. I-5. RAFINESQUE, C. S., Odatelia, new genus of North American shells (Atlantic Journal and Friend of Knowledge, 1832, p. 154). Poutson, C. A., A monograph of the Bivalve shells of the Ohio, 1832 (Translated from Rafinesque, ’20). Occasional Papers of the Museum of Zoology 73 1834° 1834-1835 1834 1835 1835-1840 1836 1838 1840 1840-1841 1842 1843 1843 1845 1846-1849 1848 Conrap, T. A., Description of some new species of fresh water shells from Alabama, Tennessee, etc. (Amer. Journ. Sci., XXV, 1834, pp. 338-343). Conrap, T. A., New Fresh-water Shells of the United States, 1834 (May); Appendix, 1835. Lea, I., Observations on the Naiades, and descriptions of new species of that and other families (Trans. Amer. Phil. Soc., V, 1834 (Aug. or Sept.), pp. 23-119). Frrussac, A. E. pg, Observations, etc., sur la Synonymie des Coquilles Bivalves de L’Amerique Septentrionale (Mag. de Zool., 1835, pp. 1-36). Conran, T. A., Monography of the Family Unionide. SCHLUETER, F., Verzeichnis meiner Conchylien. Lea, I., Description of new fresh water and land shells (Trans. Amer. Phil. Soc., VI, 1838, pp. 1-154). Swatnson, W., A Treatise on Malacology, London, 1840. Lea, I., Descriptions of New Fresh Water and Land Shells (Proc. Amer. Phil. Soc., I, 1840, pp. 284-289). Continuation (ibid., II, 1841, pp. 30-34, 81-83). Lea, I., Description of New Fresh-water and Land Shells (Trans. Amer. Philos. Soc., VIII, 1842, pp. 163-252). Lea, I., Description of Twelve New Species of Uniones (Proc Amer. Phil. Soc, 1V, 1843: p. 11). DeKay, J. E., Zoology of New York, V, Mollusca. Lea, L., Descriptions of New Fresh-water and Land Shells (Proc. Amer. Phil. Soc., IV, 1845, pp. 162-168). HERRMANNSEN, A. N., Indicis Generum Malacozoorum Pri- mordia, 1846-1849; Supplement, 1852. Lea, I, Description of New Fresh-water and Land Shells (Trans. Amer. Phil. Soc., X, 1848, pp. 67-101). Acassiz, L., Uber die Gattungen unter den nordamerikan- ischen Najaden (Arch. ftir Naturgesch., XVIII, 1852, pp. 41-50). Lea, I., Descriptions of New Species of the Family Union- idae (Trans. Amer. Phil. Soc., X, 1852, pp. 253-204). Conrap, T. A., Synopsis of the Family of Naiades of North America (Proc. Acad. Phila., VI, 1853). Lea, I., Description of Twenty-seven New Uniones from Georgia (Proc. Acad. Phil., IX, 1857, pp. 169-171). Lea, I., Description of New Unios from Tennessee, etc. (Proc. Acad. Phila., 1858, pp. 40-41). ApAms, H. and A., The Genera of Recent Mollusca, II, pp. 489-509. 74 1861 1864 1865 1868 1803 1896 1808 1900" 1900° 1900 190! IQOI 1910 IQI0* 1910” IQII IQI2 1914 1914° 1914¢ 1914 University of Michigan Lea, L, Descriptions of Twenty-five New Unionide from Georgia, Alabama, Mississippi, ‘Tennessee and Florida (Proc. Acad. Phila: Vj) 1861," p) 3848): Reeve, L., Conchologia Iconica, XVI, Unio. Lea, I., Descriptions of Eight New Unios from the United States (Proc. Acad. Phila., IX, 1865, pp. 88-89). Lea, IL, Description of Sixteen New Unionide (Proc. Acad. Phila., XII, 1868, pp. 143-145). FiscHer, P., Aanp Crossr, H., Mission Scientifique au Mex- ique et dans l’Amerique Centrale, Pt. 7. Piuurspry, H. A., ann RuHoAps, S. N., Contributions to the Zoology of Tennessee, No. 4, Mollusks (Proc. Acad. Phila., 1896, pp. 487-506). Stmpson, C. T., Unionide in: Baker, F. C., The Mollusca of the Chicago area (Bull. Chicago Acad. Sci., 3, 1808, pp. 47-110). Simpson, C. T., New and Unfigured Unionide (Proc. Acad. Phila., 1900, pp. 74-86). Stmeson, C. T., Synopsis of the Naiades or Pearly Fresh- water Mussels (Proc. U. S. Nat. Mus., XXII, pp. 501-1044). Catt, R. E., A Descriptive Illustrated Catalogue of the Mollusca of Indiana (24th An. Rep. Dep. Geol. and Nat. Res. Indiana (1899), 1900, pp. 337-535). Stmpson, C. T., Alasmidonta marginata (Say), (Nautilus, XV, I90I, pp. 16-17). Fox, W. J., The locality of Say’s Type of Alasmidonta marginata (Nautilus, XV, 1901, p. 47). Vanatra, E. G., Unionide from S. E. Arkansas and N. E. Louisiana (Nautilus, XXIII, 1910, pp. 102-104). WALKER, B., On the validity of Unio undatus Barnes (Nau- tilus, XXIV, 1910, pp. 6-10, 16-25). WALKER, B., Notes on Truncilla, with a Key to the Species (Nautilus, XXIV, 1910, pp. 75-81). OrtTMANN, A. E., Monograph of the Najades of Pennsyl- vania (Mem. Carnegie Mus., IV, I91I, pp. 279-347). OrtMANN, A. E., Notes upon the Families and Genera of the Najades (Ann. Carnegie Mus., VIII, 1912, pp. 222-365). Stmpson, C. T., A Descriptive Catalogue of the Naiades or Pearly Fresh-water Mussels. Frierson, L,. S., Remarks on Classification of the Unionide (Nautilus, XXVIII, 1914, pp. 6-8). Frierson, L. §., Observations on the genus Symphynota Lea (Nautilus, XXVIII, 1914, p. 40). OrTMANN, A. E., Studies in Najades (Nautilus, XXVIII, 1914, Pp. 41-47). Occasional Papers of the Museum of Zoology a5 1915 1915 1915 1916°* 1916* 1916¢ 1916 1916¢ 1916 1017 1918" 1918 1918¢ 1919 1919 1920 1921 Frierson, L. §., Lasmigona subviridis Conrad, redivivus (Nautilus, X XIX, 1915, pp. 57-59). WALKER, B., Unio viridis Conrad (Nautilus, X XIX, 1og15, pp. 74-78). Vanatta, E. G., Rafinesque’s Types of Unio (Proc. Acad. Phila., 1915, pp. 549-559). Frierson, L. S., Observations on the Unio cor of Conrad, (Nautilus, XXIX, 1916, pp. 102-104). WALKER, B., Pleurobema lewisi (Lea) (Nautilus, X XIX, 1916, pp. I14-116). Wa ker, B., The Rafinesque-Poulson Unios (Nautilus, XXX, 1916, pp. 43-47). OrTMANN, A. E., The Anatomy of Lemiox rimosus (Raf.) (Nautilus, XXX, 1916, pp. 39-41). Frierson, L. S., Observations on Unio giganteus Barnes (Nautilus, XXX, 1916, pp. 61-64). Urrersack, W. I., The Naiades of Missouri (Amer. Mid- land Nat., IV, 1916, pp. 1-200). Frierson, L. S., New Genera and Species of Central Amer- ican Naiades (Nautilus, XXXI, 1917, pp. 47-49). WALKER, B., Notes on North American Naiades (Occ. Pap. Mus. Zool., Univ. Mich., XLIX, 1918 (March), pp. 1-6). OrTMANN, A. E., The Najades (Fresh-water Mussels) of the Upper Tennessee Drainage, with notes on synonymy and distribution (Proc. Amer. Phil. Soc., LVII, 1918 (Octo- ber), pp. 521-626). WALKER, B., A Synopsis of the Classification of the Fresh- water Mollusca of North America, North of Mexico, and a Catalogue of the more recently described Species, with notes (Univ. Mich. Mus. Misc. Publ., VI, 1918, pp. 1-213). Frierson, L. S., Remarks upon the Identity of Unio fas- ciata Rafinesque (Nautilus, XXXII, 1919, pp. 139-141). Ortmann, A. E., A Monograph of the Naiades of Penn- sylvania, Part 3, Systematic Acount of the Genera and Species (Mem. Carnegie Mus., VIII, 1919, pp. 1-384). Frierson, L. S., Lasmigona viridis Rafinesque (Nautilus, XXXIII, 1920, pp. 127-130). OrtMANN, A. E., The Anatomy of Certain Mussels from the Upper Tennessee (Nautilus, XXXIV, 1921, pp. 81-91). 4 De 4 5 ee Sek As STS aa = roy! Paine: Ye eee . « A > , ‘ Pa it io is Sis han m La, Fi Gee ty, a - Hie ieee 8” tlh Pas ws seeps 4 on “" ; ROME FANT! tad eel ff Seine ? (ap agi fi AEN, eal 4b hs Exar: 5. . ~ a a 7 P ras —_ ae “i i ‘ ou a i 7 V.. THE NAIADES OF THE GREEN RIVER DRAINAGE IN KENTUCKY, ; By ARNOLD E. ORTMANN. Reprinted from Annals of the Carnegie Museum, Vol. XVII, No.1, 1926 Issued November 6, 1926 oa: ‘ } 7 V. .THE NATADES OF THE GREEN RIVER DRAINAGE INGIGENTUCKY: By ARNOLD E. ORTMANN. (PLATE VIII, MAp.) A number of years ago I pointed out (Ortmann, 1913,' pp. 305, 308-310, 382), that the southern tributaries of the Ohio from West Virginia as far as the Licking River in Kentucky possess a Naiad- fauna, containing only such types as are found in other parts of the Ohio-drainage, and that forms, which show exclusive affinities to the Cumberlandian fauna (Ortmann, 1924, p. 40 et seq.; 1925, p. 364 et seq.) are absent, although there are many species of wide distribution, which belong to both regions. Farther westward, the rivers of the state of Kentucky were poorly known at the time I first wrote, but in view of the fact, that the Cumberland River, containing many true Cumberlandian types ( Cf. Wilson and Clark, 1914) belongs to this group of rivers, is also a tributary of the Ohio, it became desirable to ascertain the character of the faunez of the intervening rivers, and discover whether they are intermediate or transitional between the Ohioan and the Cumberlandian type, or whether there is here, somewhere, a sharp line separating these two faune. As for the Kentucky River, the question of the general character of the fauna has been solved by Danglade (1922). He enumerates forty forms? (/. c. p. 5). All of the forty are well known as belonging to the Ohio system (many of them also to the Cumberland and Tennessee), with the possible exception of one species: Alasmidonta minor (Lea). I have previously pointed this out (Ortmann, 1925, p. 344), regarding this as a Cumberlandian species, which had invaded the upper Kentucky drainage. I now have changed my opinion, and believe that A. minor is an absolute synonym of A. calceolus (Lea), a 1 The references in parentheses refer to the titles given in the Bibliography at the end of this paper. * The list given by Danglade evidently is not quite complete, since the lower reaches of the Kentucky are still poorly known. 167 168 ANNALS OF THE CARNEGIE MUSEUM. species belonging to the Ohio drainage (see below). Thus the fauna of the Kentucky River distinctly is an Ohioan fauna, and in this respect resembles the more eastern tributaries of the system (Licking, Big Sandy, etc.). Between the Kentucky and the Cumberland is Green River. We possess a list of shells chiefly of Barren River, a tributary to Green River, published by Price (1900), in which some “doubtful Unios” have been named by Simpson. This list is of little value, since it does not give exact localities, and since some identifications obviously are incorrect. In addition, a number of synonyms are quoted as distinct species. Besides this list, there are some older (Rafinesque, 1820 and 1831; and Call, 1885), and some more recent (Simpson, 1914), generally rather vague records from Green River, sometimes containing geographical mistakes. During the last few years, however, the fauna of Green River has been studied more intensively. I myself stopped off three times in this region, and was favored by fortune in being able to obtain a good representation of the Green River Naiad fauna. Further, Mr. W. J. Clench collected in this drainage during two seasons, 1924 and 1925, for the Museum of the University of Michigan, and I was granted the privilege of examining his material collected in 1925, for which I wish to express my best thanks. In addition, Mr. B. Walker of Detroit had the kindness to send me a list of Kentucky Naiades represented in his collection, in which I found a number of records from the Green River drainage. This contained also an enumeration of the Naiades collected by Clench in 1924. I am under great obliga- tions to Mr. Walker for this. The localities, at which the recent collections wefe made are the following. (They are referred to in the text as indicated by the capital letters.) Mm—Green River, Mammoth Cave, Edmonson Co., Ky., Sept. 6, LOZ te GAO) In riffles at the lower end of an island, about three-quarters of a mile above the steamboat-landing at Mammoth Cave (head of navigation). Condition of river fair, water low, but not very clear. I found living shells by feeling for them in water from one to three feet deep, but also secured a great number of dead shells, recently taken out by muskrats. OrTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. 169 —Green River, Great Onyx Cave, Edmonson Co., Ky., Sept. 9, 7922. “(A. BE. ©:) About half a mile North of Great Onyx Cave, in riffles around a small island. Conditions splendid, water low and clear, shells extremely abundant, all found alive, in gravel and strongly flowing water, one to twelve inches deep. Mf —Green River, Munfordville, Hart Co., Ky., Sept. 24, 1925. R G D (Wee CreStae 223): “South side of River, on gravel-riffle, from three hundred to five hundred feet below bridge, a few taken in mud. River quite broad at this point, one to two feet of water.” —Green River, Rio, Hart Co., Ky., 1924 and Sept. 22, 1925. (Wi Jh) Ge; Sttae-220)). “On gravel-riffle, one half mile below bridge. Water two to twelve inches deep, fairly clear, rather swift current, not cold. Many shells had just died; as the water receded it left hundreds of them in small holes or depressions in the gravel-riffle...”’ “hundreds were located in two to twelve inches of water so thick that they touched one another.”’ (Sept. 22, 1925.) —Green River, Greensburg, Green Co., Ky., Sept. 19, 1925. (Wea €s3 Stas 217.) “Six miles West of Greensburg. Collected on pebble-bar. Unionidz rare. One to two feet of water, clear, cool.” “I believe that most of the Unionide have been killed here by , pearl-hunters..’ —Green River, eight miles South of Campbellsville, Taylor Co., Ky., Sept) 10,.1025- .(W..)..©.; Sta. 293): “Riffle, five hundred feet below bridge, gravel and small stones. Water clear, moderate current, two to eighteen inches deep.” —Green River, Dunnville, Casey Co., Ky., Sept. 15, 1925. (ii @ssSta7203): “Mouth of South Fork Creek, two miles northeast of Dunn- ville. Water clear, gravel and slate bottom; rather swift, cool, one to three feet deep. Unionide not common.” —Barren River, Bowling Green, Warren Co., Ky., Aug. 11, 192400 (AZ 2O:) ““Ewings Ford,”’ about three miles west of and above Bowling 170 ANNALS OF THE CARNEGIE MUSEUM. Green. A ford and an island, surrounded by riffles, with conditions of great variety. Water low and clear, collecting fine. Shells very abundant, in shallow water not more than one foot deep, all found alive, with the exception of two forms. S —Barren River, seven miles east of Scottsville, Allen Co., Ky., Sept. 25, 8925. (GW. JxiG.. Stas 226): “Unionidae on small gravel-riffle, in two to fourteen inches of water, water clear, swift, and rather warm. The undulate form very common, other species not so abundant.” A few other localities, chiefly furnished by Walker, which will have to be referred to only once or twice, will be given in full at the proper places. ANNOTATED LIST’ OF GREEN. RIVER NATADES: Material from the Green River has been examined by myself in the case of those forms which have an asterisk prefixed. 1. Fusconaia ebenus (Lea): R. Reported by Walker. This is the only place given for this species, and there is some doubt connected with it. I have seen no specimens belonging here. It is a species of the lower parts of the Ohio, Cumber- land, and Tennessee, and it might be present also in the lower parts of Green River. *>. Fusconaia subrotunda (Lea): Mm; O;R. This probably is the Quadrula globata of Price. The real globata (Lea) is a synonym of F. pilaris (Lea), which is given for Green River by Simpson (1914, p. 894), and mentioned by Walker in the list of his collection. However, F. pilaris is only a dwarfed form of F. subrotunda from the upper Tennessee. I have four specimens from Mammoth Cave, with the diameter 54-58 pr. ct. of the length, and one specimen, a male from Great Onyx Cave, with the diameter 56 pr. ct. From Rio I have seen seven specimens, with diameter 50-59 pr. ct. The maximum length (Rio) is 89 mm., distinctly greater than in typical pilaris. I am absolutely unable to distinguish these shells from the species subrotunda of the Ohio. OrTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. iliz/al *3. Fusconaia subrotunda kirtlandiana (Lea): R. Walker has reported to me that one specimen from Rio had the following dimensions: L. 92.5; H. 66; D. 35.5 mm., which would make the diameter 38.3 pr. ct. Four additional specimens from Rio, examined by myself, have the diameter 45-49 pr. ct. This makes these specimens of kirtlandiana to agree with the headwaters-form of subrotunda in the Ohio drainage. This form turns up at, or near, the upper limit of the range of swbrotunda, and at Rio the two forms pass into each other. *4. Fusconaia flava (Rafinesque): R. Rafinesque (1820, p. 305) cites this from small tributaries of Green River; and Price lists it as Quadrula rubiginosa Lea. Walker reports it from Rio, and I have seen from the same place a single individual with the diameter 45 pr. ct., thus being a typical F. flava, which is the form peculiar to smaller rivers and creeks. *s. Fusconaia flava trigona (Lea): B;S. Also cited by Price (as Quadrula trigona Lea, ‘‘common’’). My specimens from Bowling Green, three males, have the diameter of 56-61 pr. ct., with the beaks not elevated, and thus are typical trigona (not wndata Barnes, in which the beaks are elevated). A single specimen from Scottsville has the diameter of 55 pr. ct., and thus stands just at the lower limit of fv7gona, and close to typical flava. *6. Megalonaias gigantea (Barnes): Mm; R. Given by Price (as Quadrula heros Say) from ‘Barren Co.,” that is to say, well up in Barren River. I found a young dead shell at Mammoth Cave, and Clench collected two good specimens of fair size at Rio. Generally it is a large-river-form. *7. Amblema costata Rafinesque: Mm; O; R; C; D; B;S. Walker reports it from the Sulphur Fork of Russell Creek, Adair Co. Given by Price (as Quadrula undulata Barnes) as ‘“‘common in all streams.’ Very abundant at Scottsville. 172 ANNALS OF THE CARNEGIE MUSEUM. 8. Amblema peruviana (Lamarck). Given by Price (as Quadrula plicata Say). This is a large-river- form, which very well might exist in lower Green River. Walker has it (as plicata?) from Green River. Its presence, however, should be confirmed. *9. Quadrula pustulosa (Lea): Mm; O; R; B. Also given by Price as ‘‘very common.’’ It seems to disappear, however, toward the headwaters, as is the rule elsewhere. 10. Quadrula quadrula (Rafinesque). Mentioned by Price (as Q. lachrymosa Lea). This should be ex- pected in the lower part of Green River. Walker has it from ‘‘Barren River, Green Co.,’’ but Barren River is not in this county! *11. Quadrula verrucosa (Rafinesque): Mm; 0O; Mf; R; C; D; B;S. Given by Price (as Tritigonia (sic!) verrucosa Raf.; U. tuberculatus Barnes). Rather abundant. In the lower parts of Green River it is represented by a peculiar dwarf race; farther up it is more normal in size. 12. Quadrula metanevra (Rafinesque). Given by Price. I have never seen a true metanevra, but only the next form. Yet the main species very likely exists in the lower parts of Green River. *13. Quadrula metanevra wardi (Lea): Mm; O;R. Walker has reported metanevra from Rio, but he states that his specimens rather are wardi. What I have seen from Mammoth Cave and Great Onyx Cave (two specimens) and from Rio (two specimens) all unquestionably are wardi, a form turning up toward the headwaters wherever found. " “14. Quadrula cylindrica (Say): B. Also listed by Price. This seems to be rare in the Green River drainage. ORTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. 173 *15. Cyclonaias tuberculata (Rafinesque): Mm; O; R; C; B; S. Given by Price (as Quadrula verrucosa Barnes). Rather abundant in Green River. At Bowling Green I only saw a dead specimen. Specimens from Mammoth Cave have also been received from Mr. B. S. Sanford of Hudson, O., collected in September, 1925. In those examined, the obesity varies from 51 to 57 pr. ct. *16. Cyclonaias tuberculata granifera (Lea): O. It is doubtful, whether ‘“ Unio grandiferus Lea’’ of Price, given as ” “rather common,”’ refers to this species, since it has been placed with ‘Unio,’ and not with “Quadrula.”’ Simpson (1914, p. 906) gives “Quadrula granifera pusilla Sps.” from Green River. My only speci- men, a female, would agree with this, but it is exactly like a normal granifera (diameter 65 pr. ct.), but smaller: the other characters given by Simpson do not hold good. 17. Plethobasus cooperianus (Lea). This species mentioned by Price (as Quadrula cooperiana) might be expected here, but recent investigations have not brought it to light. 18. Plethobasus cyphyus (Rafinesque). This species is aiso listed by Price (as Quadrula e@sopus Green): it should certainly be expected to occur here, but it has not been found in recent times. *19. Pleurobema cordatum (Rafinesque): Mm; O; R; B. Given by Price (as Quadrula obliqua Lamarck). It is rather abundant, but disappears upstream. *20. Pleurobema cordatum plenum (Lea): O. Rare, and not very typical. *21. Pleurobema cordatum catillus (Conrad): Mm; O; R; B; S. Also in Walker collection from Mammoth Cave, and mentioned by Price (as Quadrula solida Lea). Rather abundant. The diameter of 174 ANNALS OF THE CARNEGIE MUSEUM. the specimens examined is over 50 pr. ct. of the length, and the nacre frequently is salmon-color or pink. *22. Pleurobema cordatum coccineum (Conrad): R; G. Given by Price (as Quadrula coccinea), and represented in the Walker collection from Greensburg, but not among the shells collected by Clench at this place. I have seen two specimens from Rio, with the diameter 47 and 49 pr. ct., thus standing close to catillus. Prob- ably more abundant in smaller streams and headwaters. *23. Pleurobema cordatum pyramidatum (Lea): Mm; O. In the Walker collection from Green River, Woodbury, Butler Co. Also listed by Price (as Quadrula pyramidata). It is not rare where I found it, with red or white nacre, and very typical in shape. *o4. Pleurobema clava (Lamarck): O; B. Also reported by Price. JI found only a few specimens, but they were very typical. *25. Elliptio crassidens (Lamarck): Mm; R. In Price’s list, “‘ Unio grandiferus Lea’ stands where this species should be expected, but it is hardly possible that it was intended (see above, under Cyclonaias tuberculata granifera). This species is rare at Mammoth Cave, as well as at Rio, and has not been found else- where. *26. Elliptio dilatatus (Rafinesque): Mm; O; Mf; R; C; D; B; S. Also recorded by Price (as Unio gibbosus Barnes). Simpson (1914, p- 600) gives Unio gibbosus armathwaitensis Wright from ‘‘Mammoth Cave, Green Co., Ky.,’’ but Mammoth Cave is not in Green Co. U. armathwaitensis is an entirely superfluous name, and an absolute synonym of dilatatus.* This species is common in Green and Barren 3 The Carnegie Museum has topotypes of armathwaitensis from ‘Branch of South Fork Cumberland, Armathwait, Fentress Co., Tenn.’ (probably Clear Fork, above Rugby), and many specimens from another stream in this vicinity, New River, Scott Co., Tenn. (collected by myself Aug. 30, 1924). All these are normal dilatatus, and the diagnostic characters given for armathwaitensis (‘shell narrower in front, widest behind, subcompressed, subsolid’’) are not at all evident. In fact, these are individual characters, which may be found anywhere among typical specimens of dilatatus. ORTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. eee Rivers, and represents a rather small race, but normal in all other respects. However, the color of the nacre varies a good deal. It may be purplish, lighter, or darker, and is very often salmon-pink or whitish. *o7. Lastena lata (Rafinesque): R. Three specimens of this rare species have been collected by W. J. Clench. 28. Arcidens confragosus (Say). In the Walker collection from Pond River, a tributary of the lower part of Green River. This is a species found in sluggish and muddy waters, and is to be expected in the lower drainage of Green River. *29. Lasmigona costata (Rafinesque): Mm; O; R; C; D; B;S. Common, given by Price as Alasmodonta rugosa Barnes. 30. Anodonta imbecillis Say. Reported by Price as from “rivers and ponds near rivers.’’ It is to be expected here, and is in the Walker collection from Bowling Green. 31. Anodonta grandis Say. Listed by Price as A. grandis and A. grandis gigantea Lea. The latter is only the pond-form of the former. This species seems to exist in this region, since Walker has both forms in his collection from Bowling Green. *32,. Anodonta suborbiculata Say: Pond near Bowling Green. The Carnegie Museum possesses four fine specimens from this locality, donated by Walker (from the Daniels collection, collected in August, 1899). 33. Anodontoides ferussaclanus (Lea). Given in Price’s list. This species might occur here, although it has not been found recently. Its re-discovery is very desirable. *34. Alasmidonta calceolus. (Lea): S. Three specimens collected by W. J. Clench. Price gives this as Alasmodonta deltoidea Lea, and in addition there is Alasmodonta 176 ANNALS OF THE CARNEGIE MUSEUM. minor Lea in his list, from Gasper Creek, Warren Co. I consider these forms absolutely identical, and the specimens from Scottsville, which I have seen, do not differ at all from specimens from the upper Kentucky drainage; and furthermore there are no essential differences from the true calceolus as found in the northern tributaries of the Ohio, nor from A. minor belonging to the Cumberland and Tennessee drainages. Simpson (1914, p. 499) suggests that A. minor may be only a local race of calceolus. In the color of the epidermis there are slight differences; typical calceolus has an ashy green epidermis and green rays, while the form of the Cumberland region is greenish yellow in ground-color, with dark green rays. Yet I have specimens from this latter region, which are exactly like calceolus in color, while darker specimens occasionally are found also in the North. The specimens from Green River and Kentucky River stand nearer to the minor-type, but otherwise there are no differences whatever to be observed. It goes without saying that it is not likely that two “‘species’’ of this type should exist in the Green River drainage. *35. Alasmidonta marginata (Say): Mf; R. Listed by Price as Alasmodonta truncata Wright. Rafinesque (1831, p. 4) nearly a century ago, cited this species from Green River as Alasmodon (Decurambis) scriptum. It seems to be rare. *36. Strophitus rugosus (Swainson): O; Mf; R; D; B;S. Reported from Rio by Walker. Given also by Price as Str. edentulus Say. It seems to be rather rare, for I collected only a few specimens, and so did W. J. Clench. Young specimens have a light brownish olive epidermis, older ones are blackish. le Ptychobranchus fasciolare (Rafinesque): Mm;0;Mf;R;C;D;B;S. Also reported by Price as Pt. phaseolus Hildreth. Specimens collected by myself at the lower stations (Mm; O; B) are all rather small, yet they often have the “humped” shape, which originated the name camelus Lea. At the upper stations, the species is quite abundant, and attains almost gigantic proportions. 2 *38. Obliquaria reflexa Rafinesque: B. “Common” according to Price. I found only a single dead shell. OrRTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. did *29. Cyprogenia irrorata (Lea): Mm; O; Mf; R; B. Mentioned by Price as ‘‘common,’’ and, where found, mostly present in good numbers. Simpson (1914, p. 328) gives C. trrorata pusilla Sps. from Green River, and, indeed, the specimens seen by me are rather below the average size. But I should hardly think that this justifies the creation of a new variety. *40. Obovaria retusa (Lamarck): O. Not rare at Great Onyx Cave, and typically developed, but not as large as in the Ohio River. 41. Obovaria subrotunda (Rafinesque): R. Given by Price (as Ob. circulus Lea). Walker reports this from Rio, and has kindly furnished measurements of four of his specimens, three of which have the diameter 60 (<), 61 (2), and 73 (c) pr. ct. of the length, and they thus belong here. 42. Obovaria subrotunda lens (Lea): R. Price lists this as Ob. lens. One of the specimens, of which Walker has given the measurements, a female, belongs here with the diameter 56) prs (Ct. *43. Actinonaias carinata (Barnes): Mm; 0O; Mf; R; G; C; D; B;S. Walker reports this from Bowling Green as var. gibba Simpson, and Price gives it as Lampsilis ligamentina Lamarck, ‘‘very common,” and L. ligamentina var. It is an abundant species, generally the prevailing one, and somewhat variable in shape. At the lower stations it usually is rather small, but farther upstream it reaches a fair size. The majority of the specimens have the typical shape of the “northern Muckett,” that is to say, that of the Ohio-form. Yet there are individuals, which approach the Cumberlandian variety called gzbba. But the same may be occasionally observed in the Ohio River, and the ‘‘southern Muckett’’ (var. gibba) cannot be credited to Green River. *44. Truncilla truncata Rafinesque: O; R. Also given by Price as Plagiola elegans Lea. 178 ANNALS OF THE CARNEGIE MUSEUM. 45. Truncilla donaciformis (Lea). Given by Price as Plagiola donaciformis. It probably exists here, since it is generally found associated with the preceding species. 46. Plagiola lineolata (Rafinesque). Given by Price (as Pl. securis Lea). It probably is found here, in the larger rivers, but has not been discovered in recent times. *47. Leptodea fragilis (Rafinesque): Mm; O; R; B. Given by Price, as Lampsilis gracilis Barnes. It appears to be common, at least at the lower stations. *48. Proptera alata (Say): Mm; O; R; B. In the Carnegie Museum, also from a pond near Bowling Green, donated by B. Walker. Reported by Price as Lampsilis alatus, ‘“‘common’’; and it may be so in the lower parts of Green River. However, it does not go far in the upstream direction. 49. Carunculina parva (Barnes). Given by Price as Lampsilis parvus. It probably exists in this drainage. 50. Carunculina glans (Lea). Reported by Call (1885, p. 31) from Green River. It may occur here, but its presence should be confirmed. 51. Micromya fabalis (Lea). Reported by Price as M. lapillus Lea. This species may belong to this drainage, since it is found elsewhere in small streams of the upper Ohio-system, but confirmation of the fact is desirable. *52. Micromya nebulosa (Conrad): D. Of forms belonging to the nebulosa-iris-group the following have been listed by Price: Lampsilis cumberlandicus Lea, L. obscurus OrRTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. 179 Lea’, L. regularis Lea, L. iris Lea, L. planicostatus Lea, L. fatuus Lea. To these Simpson, (1914, p. 119), adds: JL. nebulosa (Conr.) from “Green River,’’ and (J. c. p. 123): JL. tenera (Lea) from Bowling Green. All these, with the exception of iris, have been recognized as synonyms, for which the oldest name is Micromya nebulosa (Conrad). Yet it seems to me, that zris is also merely a form of this species. It is distinguished from nebulosa by the character of the rays, which are fine, not interrupted, and not very distinct. But M. nebulosa of the Tennessee and Cumberland drainages is extremely variable in the color-pattern. It mostly has broad, distinct, and often inter- rupted rays, which, however, frequently may be missing. Specimens with the iris-pattern do exist in the upper Tennessee, and especially in the Cumberland. On the other hand, the typical zrzs of the upper Ohio drainage never shows the nebulosa-pattern. But then again in the Lake-region, the supposed variety of zris, called novi-eboraci Lea, distinctly has it (interrupted, distinct rays), and it is impossible for me, to distinguish novi-eboraci from nebulosa, when I do not know the locality. The Cumberlandian nebulosa is also variable in the color of the nacre (whitish, salmon, or purplish), while zr7s and novi-eboract are always white inside. From Green River at Dunnville I have examined two specimens collected by W. J. Clench. One of these is fairly a nebulosa, with yellow-brown epidermis, and rather broad, distant, blackish green rays, practically uninterrupted, stronger on the posterior part of the shell, but present and distinct also on the anterior. The other speci- men is more like zr7s, with yellowish green epidermis, and fine, dark green rays, not very strongly marked, rather crowded on the posterior part, more distant on the anterior part. The nacre in both specimens is white. This tends to show, that mnebulosa and iris are conspecific (the anatomy of the two forms is practically identical), but, of course, the material at hand is too meagre to finally settle the question. Addi- tional material is to be looked for in small tributaries of Green River. The present specimens come from the uppermost station, and it is a general rule elsewhere that the nebulosa-iris forms prefer the small streams and headwaters. 4 Given on the same line with L. ovatus, but probably without the intention of making them synonyms, which would be ridiculous. 180 ANNALS OF THE CARNEGIE MUSEUM. I call my specimens nebulosa with the distinct understanding, that one of them should rather be called iris. If it should prove to be correct, that nebulosa and iris are the same, nebulosa should be stricken off the list of the Cumberlandian types (Ortmann, 1924, p. 42). It is a species of very wide distribution, going northward as far as the Lake-basin, and exclusively represented in the upper Ohio drainage by the color-phase of zris. *53. Micromya lienosa (Conrad): R; D. Shells reported by Price as Lampsilis lienosus*®, L. caliginosus Conrad, and L. nigerrimus Lea probably belong here. Walker gives lienosa from Mammoth Cave and Bowling Green. At both these places I only collected the next species (ortmanni). In addition, Walker has M. vanuxemensis (Lea) from Gasper Creek and from Rio. Sketches of the latter specimens (Rio), submitted to me, show, however, that they resemble /ienosa, since according to Walker they have purplish or white nacre, and not the peculiar salmon-tint of ortmanni. I did not see specimens of /ienosa among the material from Rio collected by Clench in 1925. But there are seven specimens from Dunnville, three males, four females according to shape. These distinctly differ from ortmannt. The shell is not so thick, the epidermis is blackish brown, rays are hardly present (only barely indicated in some). The nacre is whitish, with no salmon-color whatever, but in some specimens with purplish tints posteriorly. The constriction of the female shell is absent, or there is only a faint trace of it. These specimens corre- spond to the white-nacred phase of lienosa, often called nigerrima Lea, as found in the western and northern extension of the range from Louisiana and Arkansas into Illinois and Indiana. *54. Micromya ortmanni Walker: Mm; O; Mf; R; B. Also in Sulphur Fork of Russell Creek, Adair Co., Walker. The type-locality is Mammoth Cave (Walker, 1925, p. 1), and I have a dead female from this place. Very likely this new species has also been collected by Price, and the form mentioned as L. vanuxe- mensts Lea is this; but some of the names quoted above under lienosa may also belong here. ®» L. lienosus stands in Price’s list on the same line with L. luteolus: in analogy to Tritigonia verrucosa Rafinesque and U. tuberculatus Barnes, this might be taken for an indication of their synonymy. But this would be preposterous. ORTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. 181 This apparently is a local type of Green River, developed out of the lienosa-stock, corresponding to a degree to the vanuxemensis- type of the Cumberlandian fauna, without being directly connected with it. 55. Ligumia subrostrata (Say). Given by Price as Lampsilis subrostratus. This may be present in the lower part of Green River. *56. Ligumia recta latissima (Rafinesque): Mm; R; C; B. Given also by Price as Lampsilis rectus Lamarck. It is present, but is not abundant. 57. Lampsilis anodontoides (Lea). Given by Price from Green and Barren Rivers, but I have not seen the true anodontoides, while I found the var. fallaciosa. The form of sand- and gravel-bottom in flowing water should, however, be ex- pected here. *58. Lampsilis anodontoides fallaciosa (Smith): R; B. Walker reports this from Rio, and I have found a single individual, a male, at Bowling Green, in mud close to the bank in quiet water above the riffles. *59. Lampsiis siliquoidea (Barnes): Mm; R; C; D. Also from Bowling Green.® Price cites this species as L. luteolus Lamarck. It generally prefers smaller streams. *60. Lampsilis ovata (Say): Mm; O; R; B. Also given by Price. It is not rare at the stations mentioned, but disappears in the upstream direction. *o1. Lampsilis ovata ventricosa (Barnes); Mm; O7 Ri GD aibaoe ae Also reported by Price as L. ventricosus from ‘‘Barren River,’’ and it is in the Walker collection from the lower Green River, Livermore, 6 A fine specimen donated by Walker originally was labeled: ‘‘Cumberland River, Bowling Green, Ky.,’’ but Bowling Green is not on the Cumberland, and this species is not found in the Cumberland drainage. Walker lists it from Bowling Green. 182 ANNALS OF THE CARNEGIE MUSEUM. McLean Co., Ky. At the lower stations it is found associated and intergrading with typical ovata, but at the upper stations it is not accompanied by the latter. *62. Lampsilis fasciola Rafinesque: O; C; B. Recorded by Price as L. multiradiatus Lea. Present in the Walker collection from Bowling Green. Not rare at the two places where I found it. *63. Dysnomia triquetra (Rafinesque): O; R; G; C; D; B. Listed by Price as Truncilla triquetra. Not abundant, but ap- parently well distributed over the system. 64. Dysnomia torulosa (Rafinesque). Given by Price as Truncilla perplexa Lea. It would be quite im- portant to have this record verified, for exact localities for this species are very few. They belong to the lower Ohio, Tennessee, and Cumber- land.? The presence of the var. gubernaculum in the Green River drainage also points to the probability that the typical form of torulosa exists here. *65. Dysnomia torulosa gubernaculum (Reeve): Mm; C; B. In the Walker collection it is represented from Green River, Greens- burg, Green Co. This apparently is the Truncilia perplexa rangiana Lea of Price. The best set (6 o'o’, 4 @ @) is that collected by Clench at Camp- bellsville. I found only a dead female at Mammoth Cave and a living male at Bowling Green. The females have the marsupial ex- pansion tinted dark green, and thus this is gubernaculum. This is the headwaters-form of torulosa, known hitherto only from the upper Tennessee, but it is not astonishing that the typical torulosa (an Ohio-type) has developed this form in Green River. 71 have not been able to find any published record for this species from the Cumberland, and have commented upon this (Ortmann, 1924, p. 45 and 1925, p. 363). Yet Walker has informed me that he has specimens of torulosa from the Cumberland. According to the labels they have gone through the hands of Wetherby and Marsh, and probably were collected at Nashville by Dr. Lindsey in 1877. This seems to establish the presence of torulosa in the Cumberland River, although the information is subject to doubt, and it is remarkable that this species never again has been found in the Cumberland. ORTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. 183 66. Dysnomia flexuosa (Rafinesque). Green River is one of the original localities given by Rafinesque (1914, p. 306) for Obliquaria flexuosa. It is a very rare shell, for which few exact localities in the lower Ohio are known, and the re-discovery of this species in Green River would be very valuable. DOUBTFUL AND SPURIOUS SPECIES. Fusconaia edgariana (Lea). Cited by Price as Pleurobema ed- gariana. This probably is a misidentification, since this species belongs exclusively to the Tennessee River, and is missing in the Cumberland as well as the whole Ohio drainage. Probably young specimens of Pleurobema cordatum have been taken for it. “Unio grandiferus Lea.”’ See above under Cyclonaias tuberculata granifera and Elliptio crassidens. We do not know what this stands for. Ptychobranchus subtentum (Say). Unio subtentus has been reported by Call (1885, p. 51) for ‘““Green and Salt Rivers, Ky.’’ This species belongs to the Tennessee and Cumberland, and is unknown in the Ohio system. Since Price does not indicate its presence, Call’s record probably is a mistake. Actinonaias pectorosa (Conrad). Cited by Price as Lampsilis perdix Lea on the same line with L. zris. Whether it is thus indicated that the two are synonyms is not clear; but, if so, it is wrong. This species (perdix = pectorosa) is a Cumberlandian shell, and has never been found anywhere else. Carunculina texasensis (Lea). Given by Price as Lampsilis texa- sensis. This is a southern species, probably not found in the Green River system, and recorded by mistake. GENERAL REMARKS ON THE GREEN RIVER NAIAD FAUNA. Among the sixty-six forms, more or less positively recognized as members of the Green River Naiad Fauna, there are no Cumberlandian types. Some, indeed, have been recorded which have been hitherto regarded as such; but, as has been indicated above, our views with regard to these should be modified. This refers to the following three cases. Alasmidonta calceolus (Lea). I have taken the form found in 184 ANNALS OF THE CARNEGIE MUSEUM. Kentucky River for A. minor (Lea) (Ortman, 1924, pp. 24, 42, and 1925, p. 345) a Cumberlandian type. The Green River form is absolutely identical with it, but it is also identical with A. calceolus of the Ohio drainage. This removes this species from the list of the purely Cumberlandian shells. Micromya nebulosa (Conrad). I have considered this a Cumber- landian shell (/. c. 1924, pp. 30, 42, and 1925, p. 355). I now think, that this does not essentially differ from M. iris (Lea) of the upper Ohio, and it is even found in a form, impossible to distinguish, in the Lake-drainage (novt-eboract). On the other hand the ir7s-type is also found in the Cumberland, and occasionally in the Tennessee, and all this forces us to remove this whole association of forms from the Cumberlandian types. Dysnomia torulosa gubernaculum (Reeve). This hitherto has been known only from the upper Tennessee. But since it is only a form derived from D. torulosa, which is an Ohioan shell, it is not strange that it turns up again in Green River, having here the same relation to the main species. This cancels gubernaculum in the list of the Cumberlandian types. In this connection it is well to point out that a case similar to the last one has come to light in Micromya ortmanni. This has a certain similarity to the Cumberlandian M. vanuxemensis. It seems to me, however, that both vanuxemensis and ortmanni are derived from the lienosa-stock of the Gulf Coastal Plain, Mississippi Embayment, and lower Ohio region. They are parallel forms, originated independently of each other under similar conditions. The only difference from the case of Dysnomia torulosa gubernaculum is that in the latter the two forms cannot be distinguished, while M. vanuxemensis and ortmanni are easily told apart. The above considerations substantiate the conclusion, that there are no characteristic Cumberlandian types in Green River. There are no shells hitherto known to occur only in the Tennessee and Cumber- land’, which have extended their range into this system, without having reached other parts of the Ohio drainage. This seems to prove that there must have been at some time in the past a disconnection of the waters of Green River and of that system in which the Cumberland fauna originated. The present connection of 8 Of course, we here disregard the fact that some of the Cumberlandian types turn up again in the Ozarks; and that others are found in the Alabama drainage. OrRTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. 185 Green River and Cumberland River, by way of the Ohio, was non- existent. In more recent times, when the present drainage features had been established, the situation became different, and an interchange of forms could take place; yet a number of them never took advantage of this opportunity. It seems that chiefly forms of smaller streams belong to this latter class, as is entirely natural. In order to bring out more clearly the fact, that there is a great contrast between the Cumberland and Green River faunas, it is well to give here a list of true Cumberlandian elements, present in the Cumberland drainage, and absent in Green River (see: Wilson and Clark, 1914; Ortmann, 1924 and 1925). 1. Fusconaia barnesiana (Lea) *12. Micromya vanuxemensis (Lea) 2. Quadrula intermedia (Conrad) 13. Dysnomia arcaeformis (Lea) *3. Pleurobema oviforme (Conrad) 14. Dysnomia brevidens (Lea) 4. Pegias fabula (Lea) 15. Dysnomia lenior (Lea) 5. Plychobranchus subtentum (Say) *16. Dysnomia haysiana (Lea) 6. Dromus dromas (Lea) *17. Dysnomia lewisi (Walker) 7. Actinonaias pectorosa (Conrad) 18. Dysnomia biemarginata (Lea) *8. Carunculina moesta (Lea) 19. Dysnomia turgidula (Lea) 9. Medionidus conradicus (Lea) 20. Dysnomia florentina (Lea) 10. Micromya trabalis (Conrad) 21. Dysnomia capsaeformis (Lea) 11. Micromya taeniata (Conrad) The species marked with an asterisk (*) have representative forms in the Ohio drainage. The place of Pleurobema oviforme is taken by Pl. clava®; that of Carunculina mesta by C. glans; that of Micromya vanuxemensis by M. ortmanni; that of Dysnomia haysiana by D. sulcata (Lea); that of Dysnomia lewist by D. flexuosa. All these representatives have been found in Green River, with the exception of Dysnomia sulcata. An explanation of this condition requires special study, and may be similar to that indicated above in the case of Micromya vanuxemensis and ortmanni; yet these forms might in part indicate other possibilities in the development of these two faunas, since the distributional facts in these cases are not all of the same character. But there remain enough others, where the contrast is very striking, and this concerns chiefly nos. 4, 6, and 9, since here also the genera ( Pegias, Dromus, Medionidus) are absolutely missing in the interior drainage. And with regard to No. 1 (Fusconaia barnesiana) and the ® This case requires further study, see: Ortmann, 1925, p. 340. 186 ANNALS OF THE CARNEGIE MUSEUM. species of Dysnomia under nos. 13, 14, 15, 18, 19, 20, and 21, we observe that they represent types of shells also without representation in the Ohio drainage. On the other hand, we have in Green River certain Naiades, which belong to the Ohio drainage, but are missing in the systems of the Cumberland and Tennessee. They are the following: Pleurobema clava (doubtful) 5. Micromya lienosa 2. Aroidens confragosus 6. Ligumia subrostrata 3. Anodonta suborbiculata 7. Lampsilis siliquoidea 4. Carunculiana glans (see above) 8. Dysnomia flexuosa (see above) Moreover, there are Ohio-species in Green River, of which we know, that their center undoubtedly was in the interior Basin, but that they subsequently invaded the lower parts of the Cumberland or Tennessee (or both), coming evidently from the lower Ohio (compare Ortmann, 1925, p. 365). The most striking cases are the following: 1. Fusconaia ebenus (lower Cumberland and lower Tennessee) 2. Fusconaia flava and flava trigona (in Cumberland, but not in Tennessee) 3. Megalonaias gigantea (lower Cumberland and lower Tennessee) 4. Amblema peruviana (lower Cumberland only) 5. Quadrula quadrula and var. fragosa (lower Cumberland and_ lower Tennessee) 6. Actinonaias carinata (lower Tennessee only) Carunculina parva (Cumberland only) 8. Lampsilis anodontoides and var. fallaciosa (lower Cumberland and lower ~ Tennessee) (Pleurobema clava might also belong here) All this tends to show, that Green River has a Naiad-fauna. closely resembling that of the Kentucky, Licking, Big Sandy Rivers, and that of the Ohio and the interior Basin in general, without any par- ticular relationship to that of the Cumberland and Tennessee. A good number of peculiar forms existing still in the Cumberland are not found to the north of it; and there are others in Green River, which find here their southern limit. Thus it is clear, that there is a sharp line between the Cumberland andGreen Rivers in southern Kentucky, separating two apparently old faunas, the Ohioan and the Cumberlandian. There is no gradual transition between these faunas in the sense, that their elements gradually disappear, when we go over the several river-systems of this region (Cumberland, Green, Kentucky, Licking, etc.). This, however, should be the case, if the two faunas had been all the time connected in one major drainage-system, as they are now. OrTMANN: NAIADES OF THE GREEN RIVER DRAINAGE. 187 The two systems, at some time in the past, were separated, the Ohioan (or whatever was its master-stream) having no connection with the Cumberlandian River (Cumberland + Tennessee). Later on, how- ever, the present conditions were established, very probably by the deflection of the Tennessee and Cumberland toward the North and toward the Ohio, and there is no question, that the northward flowing parts of these rivers are of rather modern origin. This union with the Ohio must have brought about a partial mingling of the old faunas, and we have introduced above evidence for the invasion of Ohioan types into the lower Cumberland and Tennessee (Ortmann, 1925, p. 365). But, of course, an exchange should have gone on also in the opposite direction, Cumberlandian elements invading the lower Ohio. I have alluded to this previously (Ortmann, 1925, p. 364, footnote; p. 370), and instances of this will be found among those forms, which are uniformly distributed over the Interior Basin and the Cumberland- Tennessee drainage. But the distribution of these forms in the Interior Basin must be studied more closely, before we can point them out. These forms indicate the present unity of these river systems, and have largely obliterated the past condition of faunal separation, prevailing at an earlier period; yet distinct evidence of the latter still remains, as we have seen above. There is yet in Kentucky the Salt River system, between the Green and Kentucky Rivers. A few scattered records from it are at hand, but no intensive collecting has ever been done there. However, it is to be expected, that this drainage also has a fauna similar to those of the Kentucky and Green Rivers, that is to say, an Ohioan fauna, without typical Cumberlandian elements. BIBLIOGRAPHY. Catt, R. E. A Geographical Catalogue of the Unionide of the Mississippi Valley (Bull. Des Moines Acad. Sci., I, 1885, pp. 5-57). DANGLADE, E. The Kentucky River and its Mussel Resources (Bur. Fisher.; Doc. No. 934. 1922, pp. 1-8). OrTMANN, A. E. The Alleghenian Divide and its Influence upon the Freshwater Fauna (Proc. Amer. Philos. Soc., LIT, 1913, pp. 289- 390). 18S ANNALS OF THE CARNEGIE MUSEUM. ORTMANN, A. E. The Naiad-fauna of Duck River in Tennessee (Amer. Midland Naturalist, IX, 1924, pp. 3-47). ORTMANN, A. E. Fhe Naiad-fauna of the Tennessee River System below Walden Gorge (Amer. Midland Naturalist, IX, 1925, pp. 321-372). PricE, S. F. Mollusca of southern Kentucky (Nautilus XIV, 1900, PP- 75-79). RAFINESQUE, C. S.. Monographie des Coquilles Bivalves et Fluvia- tiles de la Riviére Ohio (Ann. Sci. Phys., Bruxelles. 1820, pp. 287— BBD). RAFINESQUE, C. S. Continuation of a Monograph of the Bivalve shells of the River Ohio. 1831, pp. I-5. Simpson, C. T. A Descriptive Catalogue of the Naiades, or Pearly Fresh-water Mussels. Detroit, 1914. WALKER, B. A new Species of Micromya (Occas. Pap. Mus. Zool. Univ. Mich., No. 163. 1925, pp. 1-6). Witson, C. B. & CLaRK, H. W. The Mussels of the Cumberland River and its Tributaries (Bur. Fisher., Doc., No. 781, 1914, pp. 1-63). : Nv \ : Jay 3 et y Be ees. gy SOC “Snej ypow ay “os Cb). pac vl] s : > 1 : Wand NEEL) 7 we ISsay 2, ites — = 4 Faerie = Ss ay4 Las |p aqdueg Rcodeas We 5 by zs e no = = . Dp rae? ite e ( ye , XS gr? 8 = RAIN LNA NI ADVNIVUG UAAIY NAD ‘ITIA 93?Id ‘TIAX ‘19A WAASOW FAIOANUVO STVNNV y i" bey ' its ae ay Pn a NS te Orin eve. ee Loe f pa 5 ‘e x a Can Ne a: SAS ed ee : few Lee Vite ia r ye j iv o ie eA n 1 f ve v8 noe ‘ Weerhry SMITHSONIAN HAI i ab OU | 3 Sn 00596 0653 ii tf LA a dy Sinha { H i hat ipesescbed bhi haa ‘ie Re eas ts Uitte ret 4 i th ICAI aT ‘ th Mire sia sh} +4 see Mn! ah u Miata ; Aig oO tape Mf Wi bay pee bre 1h Neat fi } dette se Pp Pate ty ba iy Beit inte bod The SRIETH pan ea at atta bebe i ty ith oe Pare th ' t Hye ti Hib | tee iy eu Hi Hy, AY ui saiaiaiuer ant a} ii warily ert Pipe!