|
1^ mj |
0) ^ li) J |
||
|
'MM.-' |
O ;> :J> r 1 |
«
)
MEMOIKS
OF THE
OAENEGIE MUSEUM
VOL. VIII
(1919-1921)
PITTSBURGH
Published by the Authority of the Board of Trustees of the
CARNEGIE INSTITUTE
PRESS OF
THE NEW ERA PRINTING COMPANY LANCASTER, PA.
/
PREFATORY NOTE
The three Memoirs bound up in this volume were issued at the following dates : No. 1, December 17, 1919; No. 2, on February 24, 1921, although it came off of the press in December, 1920; and No. 3, on June 24, 1921. The delay in the issue of the second Memoir was due to causes over which the Editor had no control, and must be charged to the irregularity existing at that time in the railway service. Though the work had been printed and the separates bound in paper and shipped, there was nearly two months’ delay in their delivery.
The first Memoir in this volume is from the pen of Dr. Arnold E. Ortmann, whose painstaking studies upon the Naiades of Pennsylvania, and especially upon those species which exist in the tributaries of the Ohio and Mississippi, have won for him a commanding position in this field of research. The collections made by him in the drainages of the Ohio and the various other eastern affluents of the Mississippi are undoubtedly the most complete now extant in any Museum. He also thor- oughly explored all the streams of Pennsylvania belonging to the Atlantic watershed and those farther south including the Roanoke River. In the preparation of this Memoir Dr. Ortmann also consulted the typical collections, which are preserved in the Academy of Natural Sciences in Philadelphia and in the National Museum in Washington, in which are the types of the species named by Say, Rafinesque, Conrad, and Lea. Through the obliging kindness of Dr. Bryant Walker, and Mr. L. S. Frierson, Dr. Ortmann had access to authentic material representing species named and determined by these authors.
As I have elsewhere taken occasion to point out, the exploration of the streams of Pennsylvania and the adjacent States occurred ‘^at the eleventh hour.” So great and so rapid has been the pollution of the streams of the Atlantic seaboard and of the rivers of the middle west, that, had not Dr. Ortmann set out fifteen years ago to determine the molluscan content of these streams, we would have nothing but tradition to guide us. It is no exaggeration to say that not only have the fishes and mussel-shells in most of these streams become extinct, but in long reaches of the rivers all animal life has been totally destroyed. Throughout the bituminous coal- regions, where mine-water charged with sulphuric acid, due to the decomposition of pyrites, has been permitted to enter the streams, the waters, which once were clear
iii
IV
MEMOIRS OF THE CARNEGIE MUSEUM.
as crystal, flow blood-red, and not even an amoeba can live in it. No more horrible illustration of the ignorance and improvidence of the citizens of the United States in dealing with great economic problems could be found. The Ohio River, “La Belle RiviO’e” of the French explorers, has become a wide sewer, into which, under our benevolent, farseeing, and wise Governments, the filth and vileness of hundreds of towns and cities, the contaminating water of thousands of mines, and the sewage of tens of thousands of cesspools and barn-yards is now being discharged in a nasty mass of corruption. “At the very nick of time,” it was resolved, before it might be too late, to secure collections representing the fluviatile life of the State of Penn- sylvania. This has been done. It cannot be done again, for, where we began fifteen years ago to collect shells from the rivers, there is now not a single shell to be found. In the spring of 1906 Dr. Ortmann made a collection of shells in the waters of the Ohio River south of Neville Island, opjiosite Coraopolis, where, as shown by the explorations of Mr. Rhoads at a somewhat earlier date, mussel-shells of numerous species were extremely abundant. In the following year Dr. Ortmann took a party of scientific friends to the locality to Show them where the shells might be found and how rich the spot was in species. To his utter chagrin, not a living specimen could be secured. All that remained were dead shells, gaping wide. Since then the barge-men have excavated the sand and gravel, in which these crea- tures once lived, and even their habitat has been destroyed.
The second Memoir is from the pen of Dr. 0. E. Jennings. It has been reviewed with most favorable comment by leading students of paleobotany. It is a fine example of the manner in which a botanist and student of vegetable ecology, familiar with the extant flora, may address himself to the interpretation of an ex- tinct flora.
The third IMemoir, upon the Mussel-shells of the South American Continent, is again from the pen of Dr. A. E. Ortmann. The Carnegie Museum is fortunate in possessing without doubt the largest collection of the mussel-shells of South America preserved with the soft parts. This collection was mainly made through the labors of Mr. John D. Haseman on the occasion of the Carnegie Museum Expedition to Central South America. While the Carnegie Museum does not as yet possess specimens of all of tlie species, which in former times have been described by those who have written upon the mussel-shells of South America, and much remains to be done in this field. Dr. Ortmann has had more material representing the animals which inhabit the shells than any other author who has ever written upon this theme. One of the most interesting results of his research is the confirmation of the affinity of numerous South American forms with those of Africa and of Australia.
PREFATORY NOTE.
V
From whatever angle we approach the study either of the fauna or the flora of South America, we arrive at the confirmation of the view, which has already long been maintained, that this Southern continent must, at one time, have been conn- ected with Africa and with Australia.
W. J. Holland.
Carnegie Museum,
June 2, 1921.
TABLE OF CONTENTS
PAGE
Prefatory Note iii
Table of Contents vii
List of Figures in the Text ix
List of Plates “ xi
Species and Varieties New to Science Described in this Volume. . xiii
Errata and Corrigenda xiv
Memoir No. 1. A Monograph of the Naiades of Pennsylvania, Part III. Systematic Account of the Genera and Species.
By Arnold E. Ortmann, Ph.D., Sc.D 1-384
Memoir No. 2. Fossil Plants from the Beds of Volcanic Ash near Missoula, Western Montana. By O. E. Jennings,
Ph.D 385-450
Memoir No. 3. South American Naiades, a Contribution to the Knowledge of the Freshwater Vlussels of South
America. By Arnold E. Ortmann, Ph.D., Sc.D 451-670
Index 671-684
vii
LIST OF FIGURES IN TEXT.
Memoir No. 1.
FIGURE PAGE
1. Glacial preserve of Margaritana margaritifera in Pennsylvania 4
2. Fusconaia subrotunda; F. s. kirtlandiana 10
3. Fusconaia flava trigona; F. flava; F. f. parvula 15
4. Amblema plicata; A. p. costata 28
5. Quadrula pustulosa; Q. p. schooler aftensis; Q. quadrula; Q. verrucosa 36
6. Quadrula metanevra; Q. cylindrica 50
7. Rotundaria tuberculata; Plethobasus cooperianus; P. cyphyus 60
8. Pleurobema obliquum.; P. o. cordatum; P. o. catillus; P. o. coccineum; P. o. pauper-
culum; P. 0. rubrum 72
9. Pleurobema clava 88
10. Elliptio niger; E. niger (Indian garbage heap); E. dilatatus; E. d. sterkii 93
11. Elliptio violaceus; E. cupreus; E. fisherianus 106
12. Lasmigona viridis; L. complanata; L. costata; L. c. eriganensis 118
13. Lasmigona subviridis 122
14. Anodonta grandis; A. g. footiana; A. ohiensis 142
15. Anodonta cataracta; A. implicata 157
16. Anodontoides ferussacianus; A. f. buchanensis 169
17. Alasmidonta heterodon; A. undulata 175
18. Alasmidonta marginata; A. m. susquehannee 183
19. Alasmidonta marginata susquehannee; A. varicosa 189
20. Strophitus edentulus 199
21. Strophitus edentulus; S. undulatus 200
22. Ellipsaria fasciolaris; Obliquaria reflexa; Cyprogenia stegaria 210
23. Obovaria retusa; 0. olivaria; 0. subrotunda; 0. s. levigata 222
24. Actinonaias ligamentina 236
25. Amygdalonaias truncata; A. donaciformis; Plagiola lineolata 241
26. Paraptera fragilis; Proptera alata 250
27. Toxalasma parvum; Eurynia fabalis; E. iris; E. i. novi-eboraci 260
28. Eurynia nasuta 273
29. Eurynia nasuta; E. recta 274
30. Lampsilis luteola; L. 1. rosacea 284
31. Lampsilis radiata; L. cariosa; L. ochracea 295
IX
X
MEMOIRS OF THE- CARNEGIE MUSEUM.
32. Lampsilis ovata; L. o. ventricosa; L. o. canadensis 300
33. Lampsilis fasciola; L. orbiculata 310
34. Truncilla triquetra; T. rangiana 327
Memoir No. 2.
1. Equisetum insculptum Jennings 399
Memoir No. 3.
1. Diagram of soft parts of female of Diplodon trifidus (Lea) 455
2. Diagram of soft parts of female of Castalina nehringi Von Ihering 456
3. Diagram of soft parts of female of Anodontites patagonica ruhicunda (Lea) 458
4. Glochidia of Hyriince, enlarged fifty times , 469
(
LIST OF PLATES.
%
PLATE :
|
I. |
Margaritana, Fusconaia. |
|
11. |
Fusconaia, Amblema. |
|
III. |
Amblema, Quadrula. |
|
IV. |
Quadrula. |
|
V. |
Quadrula, Rotundaria, Plethobasus. |
|
VI. |
Plethobasus, Pleurobema. |
|
VII. |
Pleurobema. |
|
VIII. |
Elliptio. |
|
IX. |
Lasmigona. |
|
X. |
Anodonta. |
|
XI. |
Anodonta, Anodontoides, Alasmidonta. |
|
XII. |
Alismodonta, Strophitus. |
|
XIII. |
Ellipsaria, Obliquaria, Cyprogenia, Obovaria. |
|
XIV. |
Obovaria, Actinonais, Amaygdalonaias, Plagiola. |
|
XV. |
Plagiola, Paraptera, Propter a. |
|
XVI. |
Proptera, Toxolasma, Eurynia. |
|
XVII. |
Lampsilis. |
|
XVIII. |
Lampsilis. |
|
XIX. |
Lampsilis. ^ |
|
XX. |
Lampsilis. |
|
XXI. |
Lampsilis, Truncilla. |
|
XXII. |
Equisetum, Sequoia. |
|
XXIII. |
Sequoia, Sabina, Thuyopsis. |
|
XXIV. |
Sequoia, Betula, Ilex, Populus, Alnus. |
|
XXV. |
Populus Zaddachi, Alnus Hollandiana. |
|
XXVI. |
Populus Zaddachi, Populus smilacifolia. |
|
XXVII. |
Betula, Quercus. |
|
XXVIII. |
Alnus, Betula, Quercus. |
|
XXIX. |
Juglans, Cyperacites, Typha. |
|
XXX. |
Alnus Hollandiana, Alnus microdontoides. |
|
XXXI. |
Quercus, Celastrus. |
|
XXXII. |
Ficus, Ilex, Acer. |
|
XXXIII. |
Aralia, Vaccinium. |
XI
|
Xll |
MEMOIRS OF THE CARNEGIE MUSEUM. |
|
XXXIV. XXXV. XXXVI. XXXVII. XXXVIII. XXXIX. XL. XLL XLII. XLIII. XLV. XLVI. XLVII-XLVIII. |
Diplodon. Diplodon. Diplodon. Diplodon. Diplodon. Diplodon, Castalia. Prisodon, Monocondylcoa, Anodoniites. Monocondylaa, Anodoniites. Anodoniites. Anodoniites. Anatomy of Diplodon. Anatomy of Diplodon. Anatomy of Diplodon, Castalia, Monocondylcea, Anodontite, |
SPECIES AND VARIETIES NEW TO SCIENCE DESCRIBED
IN THIS VOLUME.
FOSSIL PLANTS.
Alnus Holliandiana Jennings, sp. nov., p. 413, PI. XXV, fig. 3; PI. XXIV, fig. 8; PI.
XXVIII, fig. 1; PL XXX, figs. 1, la, Type; fig. 3, cones.
Alnus microdontoides Jennings, sp. nov., p. 415, PI. XXIV, fig. 7; PI. XXX, figs. 2, 2a, Type.
Aralia longipetiolata Jennings, sp. nov., p. 424, PI. XXXIII, figs. 1, la, 2, 2a, Type. Betula multinervis Jennings, sp. nov., p. 411, PI. XXIV, fig. 4; PL XXVII, figs. 1, la, lb, Ic, 2; PL XXVIII, fig. 2.
Celastrus parvifolius Jennings, sp. nov., p. 422, PL XXXI, figs. 2, 2a, Type.
Equisetum insculptum Jennings, sp. nov., p. 398, PL XXII, fig. 2.
Ficus (?) prunifolia Jennings, sp. nov., p. 420, PL XXXII, figs. 1, la. Type.
Ilex fur ciner vis Jennings, sp. nov., p. 421, PL XXIV, fig. 5 and PL XXXII, figs. 2, 2a, 2b, Type.
Juglans pentagona Jennings, sp. nov., p. 410, PL XXIX, figs. 1, la, 2, 2a.
Quercus approximata Jennings, sp. nov., p. 419, PL XXVII, figs. 3, 3a, Type.
Quercus laurisimulans Jennings, nom. nov., p. 416, PL XXVIII, fig. 3; PL XXXI, figs. 3, 3a.
Sequoia ohlongifolia Jennings, sp. nov., p. 400, PL XXIII, figs. 1, la.
Vaccinium palmocorymhosum Jennings, sp. nov., p. 426, PL XXXIII, figs. 3, 3a, Type, and 4, 4a.
MOLLUSCA.
(Naiades.)
Alasmidonta (Decuramhis) marginata suquehannce Ortmann, var. nov., p. 187; PL XII, fig. 4; text-fig. 19.
Diplodon hasemani Ortmann, sp. nov., p. 478; PL XXXIV, figs. 1-4; PL XLVII, fig. 5; text-fig. 4a.
Diplodon imitator Ortmann, sp. nov., p. 491; PL XXXIV, figs. 5-7; PL XXXV, figs.
1-2; PL XLV, fig. 1; PL XLVII, fig. 6; text-fig. 4b.
Diplodon simillimus Ortmann, sp. nov., p. 495; PL XXXV, figs. 3-6; PL XLV, fig. 2; text-fig. 4c.
Diplodon .vicar ius Ortmann, sp. nov., p. 497; PL XXXV, figs. 7-8; PL XXXVI, figs. 1-2; PL XLV, fig. 3; text-fig. 4d.
Diplodon decipiens Ortmann, sp. nov., p. 499; PL XXXVI, figs. 3-6; PL XLV, fig. 4; PL XLVII, fig. 7; text-fig. 4e.
xiii
XIV
MEMOIRS OF THE CARNEGIE MUSEUM.
Diplodon hildce Ortmann, sp. nov., p. 514; PI. XXXVI, fig. 7; PI. XXXVII, figs. 1-2; PL XLVI, fig. 3; text-fig. 4i.
Diplodon mogymirim Ortmann, sp. nov., p. 520; PI. XXXVII, figs. 4-7; PI. XLVI, fig. 5; PI. XLVIII, fig. 2; text-fig. 4k.
Diplodon herthce Ortmann, sp. nov., p. 528; PI. XXXVIII, figs. 1-4; pi. XLVI, fig. 6. Diplodon enno Ortmann, sp. nov., p. 531; PI. XXXVII, figs. 5-8; PL XLVI, fig. 7. Monocondylcea obesa Ortmann, sp. nov., p. 583; PL XL, figs. 4-6.
M onocondylcea hollandi Ortmann, sp. nov., p. 585; PL XLI, fig. 1.
Anodontites hyrioides Ortmann, sp. nov., p. 604; PL XLII, figs. 3-5.
Anodontites hasemani Ortmann, sp. nov., p. 609; PL XLII, figs. 6-7.
ERRATA AND CORRIGENDA.
P. 106. Fig. 11, for “Elliptis” read Elliptio.
P. 418. Fifteenth line from top, for “Daphnes” read Daphnea.
P. 426. Twelfth line from bottom, for “ Vaccinurn” read Vaccinium.
Fublieations of the Carnegie Museum, Serial No. 104-
MEMOIKS
. OF THE
OAENEGIE MUSEUM.
VOL. VIII. No. 1.
W. J. HOLLAND, Editoe.
A MONOGRAPH OF THE NAIADES OP PENNSYLVANIA. PART III SYSTEMATIC ACCOUNT OP THE GENERA AND SPECIES
By AENOLD E. OETMAHE", Ph.D., So.D.
PITTSBUEGH.
Published by the Authoeity of the Boabd of Teustebs of the CAENEGIE mSTITTJTE.
Decbmbee 17, 1919.
PRICE LIST OF PUBLICATIONS OF THE CARNEGIE MUSEUM
AimUAL BEPO&TS OF THE DIBECTOB
1898, 30o. (searoe); 1899, 85o.| 1900, 30c. (scarce) | 1901-19, 2Sc. each.
BEPOBT8 OF FBOOEEDINOS OF FOONDEB’S DAT
1898-1919, 350. eack
AHNALS OF THE OABNEOIE MUSEUM
The Annals are supplied to those who subscribe in advance in parts (paper-bound), as publi^ed, @ $3.50 pm volume; Vols. I-Xn, 1901-1919, bound in green doth @ $4.00; bound in } Morocco @ $4.50.
MEMOIBS OF THE OAi^OIE MUSEUM
The Memoirs are supplied to those who subscribe in advan ce in parts (paper-bound), as published, at $10 per vblnme; bound in green buckram at $10.75, and in half-morocco at $12.00.
VOL. L (1901-3)
No. 1. Dijdodocn^ Its Osteology, Taxonomy, ete. No. 3. Ostedogy of the Steganopodes. SHmujn* $2.75
Hatcher $3.00 ** 4. dassification of Chalcid Flies. Ashuead.. 6£0
** 2. Oligocene Oanida. Hatches 1.60
VOL. n. (1901.^)
No. 1. OsteologyofHajdocanthosaums, etc. Hatches $2.00 ** 6. Osteology of D^odocns. Houahd ....... 1.60
** 2. Osteology of Baptanodon. Hmuoss 1.60 7. Ostedoinr of Pxotostega. WisnAHo ....... .76
** 3. Fossil Avian Bemaias from Annissan. East- ** 8. New Suilline Remains from Oie Miocene of
HAK 1.00 Nebraska. Pkcebson .76
** 4. New Bodents and Discussion of Origin of ** 9. Notes on the Osteology of Baptsmodon, etc.
Dnmemeliz. Petsbsoh 1.76 Oismose 1.00
No. 5. Tertiary of Montana. Dohgiass .....$1.00 10. The Crawfishes of Pennsylvania. Obthann AOO
voL.in.
No. 1. Arduedogical investigations in OOsta Bica. No. 2. Osteology of Moropus. Hoixahd and Petee-
Habtaian $6.00 SON $6.00
VOL. IV.
No. 1. Emdy Chinese Writing. Ghaupant $3.00 No. 5. New Carnivores from the Miocene of West-
** 2. Formosan Fishes. Jordan .26 em Nebraska. Petesson .....$1.60
** 3. Bntdodontidse. Petebson 2.60 ** 6. Monograph of the Najades of Pennsylvania.
** A Fishes of Formosa. Jordan and Bichabd- Pts. I and n. Obtaeann. 2L60
SON 1.25 ** 7. Catalog Eocene Fishes from Monte B<dea in
Carnegie Ifigsenm. Eastman 3.0$
VOL. V.
The Fresh Water Fishes of British Guiana. Eisenaiann. $10 unbound; $10.75 doth; $12.00 | morocco.
VOL. VL
No. 1. Fishes from Korea. Jordan and Msax .... $1.50 No. A Fishes obtained in Japan. 1911. JoimAN
** 2. Lantem-fidies of Japan. Gilbert 1.00 and Thompson ....................... $SB0
** 3. Oymnotid Eds of Tropical Amarifta., Nos. 6-7. Fossil Fishes in the Cam^e Moseinn.
Mapkh igT.T.TR, 1.60 Parts II— IV. Eastman 6.00
VIL
No. 1. The Cheirodontinse. EKSSNAtANN $3B0 No. 3. Opbidia from S. America. Grippin ....... 2.00
** 2. Turtles of Uinta Formation. Ghmore .... 2.00 No. A PirndodeDa and 'l^pblobagros. Ekenaiann. 2.00
No. 5. The Pygldlldm. Ekwaaann 2.60
BEPBINTS FBOM THE AHHALS OF THE CABNECHE MUSEUM
Nos. 1-12. See preceding lists. (Out of prmf.)
12. Skdetan of Titanothexinm di^pr. Hatcher. . $ B5
13. Fore Idnibs and Manns of Brantosaums.
Hatcher .60
lA The TtadtodontidB. Hatcher .25
15. New Pmmsg^vania Thtana Ashe .25
16. Ostedogy of FsittacL Shdnrudt B6
17. Annotated Catalogue of Genus il^irtiila. H. H.
Saiith 1.26
18. Two New Babaman B^doptean. Holland ... JLO
19. Elosaums Parvus. Petebson & Gilmobe J6
20. Boundary Contcoveesy Between Fenn^vania
and Viri^iia, 1748-1785. C^umbinb JOO
MEMOIKS
OF THE
OAENEGIE MUSEUM.
VoL. VIII. No. 1.
A MONOGRAPH OF THE NAIADES* OF PENNSYLVANIA. PART HI. SYSTEMATIC ACCOUNT OF THE GENERA AND SPECIES.'
By Arnold E. Ortmann, Ph.D., Sc.D.
(Plates I-XXI.)
Introductory.
In the first part of this monograph (Ortmann, 19116),^ certain anatomical features hitherto but little investigated were considered {1. c., Pt. I, p. 282 ff.), and their bearing upon the rearrangement of the system of the Naiades was dis- cussed in Part II (1. c., pp. 322 ff.). At the end of Part H, a key for the genera was attempted (pp. 335-338). Since then additional material, chiefly representing
* In Parts I and II of this Monograph, which appeared in the Memoirs of the Carnegie Museum, Vol. IV, No. 6, I employed the term Najades, following the usage of certain well-known authors who have written upon this group. The spelling of this word given by Lamarck, who first employed it, was Nayades, and’ this would have priority according to a strict application of the laws governing nomen- clature, were it not for the fact that the Lamarckian term plainly is an error in transcription from the original Greek, which is Na’tdSes. I, believe that when a Greek word is used it ought to be transcribed as nearly as possible in conformity with philological usage, and therefore have reverted to the form Naiades, which is not only good Greek, but sanctioned by the usage of a multitude of other authors. The chief end of the “ law of priority ” is not to preserve and perpetuate mistakes in spelling, even when made by men as eminent as Lamarck.
^ This paper is in continuation of the papers published in the Memoirs of the Carnegie Museum, Vol. IV, No. 6, 1911.
^ The references given in the text refer to the bibliography at the end of this paper,
1
2
MEMOIRS OF THE CARNEGIE MUSEUM.
such forms as are not found in Pennsylvania, has been secured and studied. This made necessary a number of changes, which chiefly affect the division into genera, and which have been published (Ortmann, 1912). The system proposed in this latter paper will be used in the following pages with such corrections as have become inevitable in the course of still further studies.
With regard to the systematic literature, I have refrained from giving a full list of references and synonyms under each species, since the citations given by Simpson (1914) are as complete as could be desired. The quotation of the earlier reports from Pennsylvania was deemed desirable, in order to get an idea of the advance made in recent years in our knowledge of the subject.
Family MARGARITANID^ Ortmann (1911).
Ortmann, 1911a, p. 129; 19116, p. 334; 1912, p. 223.
Genus Margaritana Schumacher (1817).
Ortmann, 1912, p. 230; Simpson, 1914, p. 511.
Type My a margaritifera Linnaeus.
1. Margaritana margaritifera (Linnaeus) (1758).
Margaritana margaritifera (Linnaeus) Simpson, 1914, p. 513.*
Plate I, fig. 1.
Records from Pennsylvania:
Lea, Obs. II, 1838, p. 56, and VII, I860, p. 225. As to the correctness of these records, see below. Hartman & Michener, 1874, p. 91. See also below.
Conner, 1904, p. 91 (Still Creek, Quakake).
Ortmann, 19096, p. 208.
Characters of shell: Shell large and heavy, cylindric-ovate, elongate, often arcuate when old. Anterior end rounded, posterior produced. Beaks very little elevated. Epidermis blackish or blackish brown. Nacre whitish, pinkish, or somewhat purplish, posteriorly iridescent. Pseudocardinal teeth present, laterals obsolete, generally entirely wanting. Inside of the mantle-line a number of small muscle-scars.
Size: My largest specimen (from Rene Mont) measures: L. 152 mm.; H. 67 mm.; greatest D. 49 mm. This is larger than any of the previous records. The maximum length given by Carl (1910, p. 65) for the form found in the Odenwald, Germany, is 136 mm. Israel (1910, p. 177) gives 140 mm. for the form from the Elster-drainage in Germany.
\
\
\
V' .
ORTMANN: monograph of the naiades of PENNSYLVANIA. 3
Soft parts and glocMdia (See Ortmann, 1912, p. 230, fig. 1; and 19136, p. 89). In the American form, the giochidia have not yet been observed. The fine teeth on the lower margin are present only in fully mature giochidia.
The breeding season in Pennsylvania occurs in the period from June to August (Conner, 1909, p. 112); in Europe it occurs from the middle of July to August (Harms, 1907, p. 818; Carl, 1910, p. 16).
Remarks: There is no possibility of mistaking this species, for it is sharply characterized by size (being the largest shell of the Atlantic-drainage) , shape, color, hinge, and by its peculiar station. Compared with the same species from other localities, the Pennsylvanian specimens are remarkable for their size, being the largest ever recorded. I cannot see any differences between them and specimens from New England and Newfoundland. European shells likewise perfectly agree with them. Specimens from the Pacific slope are all considerably smaller and comparatively thinner, and the color of the nacre inclines more toward purplish tints. There are also slight differences in the hinge. This western form has been distinguished as a variety (falcata Gould) .
Localities represented in the Carnegie Museum: ^
Locust Creek, Tamaqua; Cold Run, Hecla; Indian Run, Rene Mont; all in Scliuylldll Co., Pennsylvania. Westfield, Hampden Co. (Hartman collection); Amherst, Hampshire Co., Massachusetts (Holland collection).
Amy Brook, Henniker, Merrimack Co., New Hampshire (G. H. Clapp).
Cape Elizabeth, Cumberland Co. (G. H. Woods); Aroostook River, Caribou, and Little Madawaska River, New Sweden, Aroostook Co., Maine (0. 0. Nylander).
South West River, Belvoir Bay (Hare Bay), Newfoundland (G. H. Clapp).
From Europe (all received from W. Israel) :
Perl Bach, Postfelden near Falkenstein, and Perlbach, Rehau above Hof, Bavaria; Goernitzbach, Oel- snitz, Saxony; Aumabach, Rohna, Saxe-Weimar; Steinach, Sonneberg, Saxe-Weimar; Ulfenbach, Affolterbach, Hesse, Germany.
From the Pacific fiope (var. falcata Gould) :
Long Valley, Lake Co., California (Hartman collection); Chehalis River, Porter, Chehalis Co. (H. Hannibal); Seattle (P. B. Randolph), and Ravenna Park, Seattle, King Co., Washington (H. Hannibal).
Distribution and Ecology in Pennsylvania (See fig. 1) : This species has been recorded by Lea from Crum Creek, Delaware Co., and by Hartman & Michener from White Clay Creek, Chester Co. Lea also says that it goes southward only as far as middle Pennsylvania. I have expressed (19096) my doubts as to the correctness of these two localities, and must maintain them most emphatically. In White Clay Creek I have collected myself, and found only the common fauna of the Atlantic streams, and Margaritana is not found where this is present, accord- ing to my experience.
^ When not otherwise indicated, the specimens have been collected by myself.
4
MEMOIRS OF THE CARNEGIE MUSEUM.
The first to report an exact locality for this species in Pennsylvania was Conner (1904) Still Creek, at Quakake, Schuylkill Co. This creek is not named on the sheets of the U. S. Topographic Atlas, but it is, as I have ascertained, a tributary of the ipipermost Little Schuylkill, and the place Quakake is no doubt the same as Quakake Junction, not far from Tamaqua. In the same general region, near Tamaqua, this species once used to be abundant in the headwaters of the Little Schuylkill, and its metro])olis was in Locust Creek (See fig. 1) but the
Glacial Preserve of Margaritayia margaritifera in Pennsylvania.
recklessness of the pearl-hunters has nearly exterminated it. At the present time living specimens are rare in Locust Creek. The natives also report that it used to be found in Pine and Ilosensock Creeks.
This species also occurs to the South of Tamaqua, in tributaries of the Little Schu3dkill, in Cold Run, above Hecla, and in Indian Run, at Rene Mont. The latter localit}^ marks the southernmost extension of the range of the species. Other creeks in this region also may have formerl}" contained this shell, but, as for instance in the case of Panther Creek, it must have been destroyed long ago by the pollution from the coal-mines.
The headwaters of the Little Schuylkill in Schuylkill County appear to be the onl3" region in Pennsylvania where this species is found. It has never been reported from any other jiart of the state (except the spurious records mentioned above), and the efforts of myself, of Conner, and of Pilsbr^^ to locate it elsewhere have only proved its absence. The writer devoted much time to hunting for it, or to securing information about its presence, in northern Schuylkill County, in
ORTMANN: monograph of the naiades of PENNSYLVANIA.
the upper Lehigh-drainage, in the Pocono Mountains, and in the region of the North Branch of the Susquehanna, but without success.
In the upper Little Schuylkill-drainage this species lives in mountain streams with cold water (trout-streams) at an elevation of 800 feet and more above sea- level. The sources of these streams are at about 2000 feet, and the greatest number of shells is found at about 1000 to 1200 feet. These streams are rough and full of little falls and rapids. The shell prefers eddies and pools which are rather deep, with a steady and lively current, and with gravelly and sandy bottoms. Sometimes these shells are found in sandy (but not muddy) bottoms of mill-ponds in quiet water, but they probably have been washed down into these. Cold water with lively currents seems to be essential, and also shade, for I chiefly found speci- mens in places where the banks of the streams were wooded, and not where they ran through open fields. The characteristic shrub in these woods is Rhododendron maximum, and besides Alnus, Carpinus, and often Tsuga.
In Europe it has been observed that Margaritana is missing in streams which run over limestone rocks, and that it is very impatient of water which holds lime (Haas, 1910, p. 109). The same is true in this country. All the streams in Schuyl- kill County in which it lives run over sandstones and shales. Indian Run is entirely in the Devonian Clinton Shales, while Cold Run and the more northern streams are in the Lower Carboniferous Mauch Chunk Shales, the boulders in the water-courses being formed by sandstones and conglomerates of the overlying Pottsville beds.^
The headwaters of the Little Schuylkill form a perfectly isolated station for this species, about one hundred miles away from the nearest locality, which is to the North, in Rockland County, in southern New York. The Pennsylvanian area of this species is not only the most southern extension of its range in eastern North America, but it also has the peculiarity of being the only one to the South of the Terminal Moraine. Thus it may be regarded as a part of the Glacial Preserve (refugium) of this species (See Ortmann, 1913a, pp. 377 ff.). Margaritana ynar- garitifera in Pennsylvania is a fine example of a Glacial Relic.®
^ In this connection I should mention that 0. O. Nylander, who sent me specimens from the Aroos- took River, Maine, states positively that the formation is Aroostook Limestone (Silurian. Cf. Williams, H. S., & Gregory, H. E., in Bull. U. S. Geol. Surv., 165, 1900, pp. 44 and 141). This matter, however, should be investigated more carefully.
® Subfossil shells of Margaritana margaritifera have been found in Hartman’s Cave, near Strouds- burg, Monroe Co., Pa., associated with shells of Elliptio violaceus {Unio complanatus) , bones of living and extinct vertebrates, and with human implements of stone, bone, and horn (See Leidy, 1889). I have seen, in the Philadelphia Academy of Natural Sciences, a left valve taken from Hartman’s Cave, which undoubtedly is this species. This locality is within the glaciated area, and thus these remains are surely
6
MEMOIRS OF THE CARNEGIE MUSEUM.
General Distribution: This species is found, outside of North America, in northern and middle Europe, northern Asia, and Japan (Simpson, 1900, p. 677).® With regard to this, we do not need to go into detail here. Its North American distribution recently has been discussed by Walker (1910a, p. 140, pi. 2), and a careful list of localities has been given, to which we should add,, however, a few new localities represented in the Carnegie Museum (see above), and further Buck- field, Oxford Co., Maine (Allen, J. A., in Nautilus, XXV, 1912, p. 120) (See also Nylander, 1914).
I do not think that Margaritana margaritifera is of North American origin, as represented by Scharff (1912, p. 51), but I believe that it reached eastern North America by the North Atlantic land-bridge, coming from Europe (See Scharff, ibid.). I also hold the opinion of Scharff that there were other places of survival during the Glacial time, either within the drift area, or close to its eastern edge, on the former eastward extension of the continent.
Family UNIONID.® (D’Orbigny) Ortmann (1911).
Ortmann, 1911a, p. 129; 19115, p. 335; 1912, p. 223.
Subfamily UNIONINiE (Swainson) Ortmann (1910).’^
Ortmann, 1910, p. 116; 1911a, p. 129; 19115, p. 335; 1912, p. 224.
Key to the Genera.
tti. All four gills serving as marsupia. ^lantle connection between anal and supra-anal openings present, short, and deciduous.
bi. Placentse subcylindrical, rather persistent, often red. Shell smooth, wdthout sculpture upon the disk. Beak-sculpture simple, concentric, and poorly developed Fusconaia.
62. Placentae lanceolate and compressed, not very persistent, whitish, or yellowish, but not red.
postglacial. This would indicate, possibly, a very early step in the postglacial dispersal of this species, provided these shells are actually from the neighborhood of this cave, and have not been carried there by man from a long distance. At present, Margaritana is not found in this region. I have hunted for it in vain in Broadhead Creek at Henry ville (above Stroudsburg), and have received the assurance from competent persons that nothing resembling this shell is found in the trout-streams within a radius of at least fifty miles from Henryville.
® It also has been reported from Iceland, but this has been questioned. The Carnegie Museum has received from W. Israel a specimen labeled “ Reykjavik, Iceland.” After correspondence with Israel and Stejneger (in Washington), and by Stejneger with parties in Copenhagen, there remains no doubt that this specimen is to be traced back to a dealer in Copenhagen, who tried to impose upon a number of conchologists. He even claimed that the specimens in question were collected by Mrs. Israel in 1863, before she was born!
’’ This subfamily cannot be credited to v, Ihering (1901, p. 53) since he used the name in an entirely different sense.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
7
Shell with sculpture upon the disk (ridges or tubercles). Beak-sculpture concentric, double-looped or zig-zag.
Cl. Chief sculpture upon disk consisting of a few oblique, strong undulations. Beak-sculpture
concentric Amblenia.
C2. Chief sculpture on disk consisting of tubercles and warts. Beak-sculpture concentric.
double-looped, or *zig-zag Quadrula.
a 2. Only the two outer gills serving as marsupia. hi. Shell more or less tubercular upon the disk.
Cl. Beak-sculpture well-developed, consisting of numerous wavy ridges. Nacre dark purplish. Soft parts whitish. No supra-anal Rotundaria.
C2. Beak-sculpture rudimentary, consisting of a few obscure, concentric bars. Nacre light-
colored. Soft parts generally pale orange. Supra-anal present Plethobasus.
62. Shell smooth upon disk, without tubercles.
Cl. Shell rather thick and solid, short and high, sometimes drawn out posteriorly, and then distinctly cuneiform and oblique Pleurobema.
C2. Shell generally not very thick, often rather thin, not short and high, but more or less elon- gated, and not oblique Elliptio,
Genus Fusconaia Simpson (1900).
Ortmann, 1912, p. 240; Simpson, 1914, p. 865 (as section of Quadrula).
Type Unio trigonus Lea (= F. flava trigona).
Key to the Species and Varieties.
ai. Shell without a distinct posterior ridge, disk rather uniformly convex. Color brownish, or blackish brown.
bi. Diameter of shell fifty percent of length, or more F. subrotunda.
&2. Diameter of shell less than fifty percent of length F. subrotunda kirtlandiana.
02- Shell with a distinct posterior ridge, disk flattened, or even slightly concave in front of the ridge.
Color lighter, more reddish or yellowish brown.
61. Shell attaining a good size. , Growth-lines irregular.
Cl. Diameter of shell less than fifty-five percent of length F. flava.
C2. Diameter of shell fifty-five percent of length or more F. flava trigona.
62- Shell much smaller. Diameter very variable, but mostly considerable. Growth-lines very distinct and regular F. flava parvida.
Fusconaia subrotunda (Lea) (1831).*
Quadrula subrotunda (Lea) Simpson, 1914, p. 892.
Plate I, fig. 2.
Records from Pennsylvania: *
Harn, 1891, p. 137 (western Pennsylvania).
Clapp, 1895, p. 116 {U. pilaris from Allegheny County is undoubtedly this).
® This should not be called F. sintoxia (Rafinesque), although Vanatta (1915, p. 558) identifies Rafinesque’s species with it. According to the measurements given by Vanatta, the diameter of Rafines- que’s sintoxia is forty-three percent of the length; and thus it could only fall under kirtlandiana. But since the nacre is described as rose-colored, this does not seem likely since subrotunda and kirtlandiana have not rose-colored nacre.
8
MEMOIRS OF THE CARNEGIE MUSEUM.
Rhoads, 1899, p. 136, has recorded specimens of this species under U. obliquus from Coraopolis, Alle- gheny Co., and Beaver, Beaver Co.
Ortmann, 19096, p. 200.
Characters of Shell: Shell large and heavy, swollen, the diameter amounting to fifty percent of the length or more, subcircular or ovate; when old, often oblique and drawn-out at the lower posterior end. No posterior ridge. Beaks moderately }irominent. Beak-sculpture not distinctly observed, but probably weakly de- veloped and concentric. Epidermis in young specimens light brown, almost yellow, with more' or less distinct dark rays, which often aiijiear as bundles of fine lines, and frequently break up into squarish spots, and with dark concentric growth- lines. In older specimens the epidermis turns darker, brown to blackish, and becomes nearly uniform without any rays.
Hinge-teeth heavy, pseudocardinals divergent in the young, becoming very heavy, and subparallel to the laterals, in old shells. Interdentum variable, but generally very wide, and lieak-cavity very deep and compressed. Nacre always whitish.
. L. H. D. Pr.ct.
Size: Industiy, Cat. No. 61.3938a 107 mm. 81 mm. 53 mm. .50
This is the largest specimen at hand. It fairly represents the average in outline, but the diameter has been unduly lowered by the production of the posterior end.
Soft parts and Glochidia (See Ortmann, 1912, p. 244).
Breeding season: Tachytictic form, breeding in June and July (Ortmann, 1909a, ]). 101). Gravid females have been found on the following dates: June 22, 1909; June 24, 1909; July 3, 1908; July 11, 1911; July 5, 1909; July 13, 1908. Glochidia as well as eggs were found on July 5 and 13; on the other dates, eggs only.® The discharge of placentae was observed in a few cases as early as June 24 (Ortmann, 19116, p. 306), but this was in captivity, and was certainly premature.
Remarks: This is one of our heaviest and largest shells. Its external char- acters are rather indifferent, and it resembles several other species, with which it is easily confounded. In general the subcircular or oval outline, with rather evenly curved margins, and the deep, compressed beak-cavities characterize it. Pleurobema ohliquuni catillus comes very near to it, however, but this form is mostly more triangular, with the lower margin more nearly approaching a straight line, and with a more distinct lower posterior angle. It has only a rather shallow, and not a compressed, beak-cavity. Of course the characters of the
® Of the form from Elk River, West Virginia (var. leucogona Ortmann, 19136, p. 89) I found gravid females as early as May 25, 1911, and as late as July 10, 1911.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
9
soft parts are entirely different, and, when females are at hand, there is no mistake possible. The soft parts of F . subrotunda are often of the orange type, of Pleurohema obliquum never. Specimens with red nacre are always obliquuni-forms, and never subrotunda.
There is no difference whatever between the sexes in the shell of this species. The differences between it and var. kirtlandiana will be discussed under the latter.
Much confusion prevails as to this species, and it has often been misidentified, probably because originally only the young was described, and no good figures of the old shell were published. The synonymy given by Simpson seems to be correct, but I think that the following reference should be added; Unio varicosus Lea (Obs. I, 1834, pi. 11, fig. 20, Ohio River). Lea compares this with Plethobasus cyphyus (= cesopus), and Simpson (1900, p. 765) identifies it with Plethobasus cicatricosus (Say). Lea’s figure would stand very well for an old Fusconaia sub- rotunda, except for its color, which is too light, but I have several old specimens with a rather light (brown) epidermis, although not as light as in this figure. The color of Lea’s varicosus is all that agrees with Pleurobema cicatricosus. According to specimens of the latter in the Carnegie Museum, and the figures of Reeve (1864, PI. 8, fig. 31, and PI. 13, fig. 50), the character of the concentric ridges of the shell is entirely different from that in Lea’s figure. Furthermore the shape of the latter is not at all the characteristic shape of cicatricosus. The specific name varicosus would have priority (1829) over subrotundus, but cannot be used, since it is pre-occupied by U. varicosus Lamarck, 1819, now Alasmidonta varicosa. Localities in Pennsylvania, represented in the Carnegie Museum:
Ohio River, Cooks Ferry, Shippingport, Industry, Beaver Co.; Coraopolis (S. N. Rhoads) and Neville Island, Allegheny Co.
Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).
Mahoning River, Mahoningtown, Lawrence Co.
Allegheny River, Aladdin, Godfrey, .lohnetta, Kelly, Armstrong Co.
Monongahela River, Westmoreland Co. (G. A. Ehrmann), and Charleroi, Washington Co. (G. A. Ehr- mann).
Cheat River, Cheat Haven, Fayette Co.
Other localities, represented in the Carnegie Museum.
Ohio River, Toronto, Jefferson Co., Ohio; St. Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio.
Tuscarawas River, Ohio (Holland collection).
Levisa Fork of Big Sandy River, Prestonsburg, Floyd Co., Kentucky.*"
Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).**
*" Only one specimen found by myself. It has the diameter of fifty percent, and thus belongs here, but stands close to the var. kirtlandiana.
** These specimens do not differ from the general run of the form from the Ohio. In the upper
(
}
10 MEMOIRS OP THE CARNEGIE MUSEUM.
Distribution and Ecology in Pennsylvania (See fig. 2) : This species belongs in Pennsylvania to the larger rivers, the Ohio, Allegheny, and Monongahela, to the Beaver as far as the lower Mahoning, and to the lower Cheat. It has never been found in any other tributary of the upper Ohio system. It prefers heavy shingle and gravel, in a strong current, and is especially adapted to this habitat by its
■ Fusconaia subrotunda.
• Fusconaia subrotunda kirtlandiana.
subglobular shape and heavy shell. In the Allegheny it occurs as high up as Kelly in Armstrong County; above this point, the river is largely polluted, so that the upper limit of its former distribution cannot be now ascertained, but it is positively missing in the upper Allegheny above Oil City, and is not represented by its variety kirtlandiana, while the latter takes its place in French Creek. In the upper parts of the Beaver-drainage it is also gradually replaced by kirtlandiana. Particulars about its range in the Monongahela are wanting, but it possibly reached a little beyond the West Virginia line. It is found in the lower Cheat, and has also been found in the Indian Garbage heap at Point Marion, opposite the mouth of the Cheat (See Ortmann, 1909c).
General Distribution:
Type locality; Ohio (Lea).
This species is positively known only from the Ohio-drainage, ranging from Pennsylvania westward to Illinois, and is generally restricted to the Ohio River
Tennessee region this species is represented by a dwarfed race, commonly called U. pilaris Lea. Also in Elk River, West Virginia, I have discovered a dwarfed race, which I have called var. leucogona (Ort- mann, 19136, p. 89), but I doubt now the propriety of distinguishing this by a name.
ORTMANN: monograph op the naiades of PENNSYLVANIA.
11
proper and some of its larger tributaries. Of the latter outside of Pennsylvania the Tuscarawas and Scioto Rivers contain it (Sterki, 1907a); it is in the Illinois River in Illinois (Baker, 1906). Call (1896a) does not mention this species frorn Indiana, but he unites it with F. ebena (Lea), and quotes this from the Ohio and Wabash. According to Blatchley & Daniels (1903) subrotunda is plentiful in the Wabash and Tippecanoe Rivers in Indiana. The form from Elk River in West Virginia (Big Kanawha-drainage) is slightly different (var. leucogona). Simpson (1900) cites the Cumberland and Tennessee river-systems, and Wilson & Clark (1914) confirm this for the Cumberland. Similar forms are, indeed, found in these rivers, but they generally go under different names (mostly pilaris Lea). Yet I have typical subrotunda from the Tennessee in northern Alabama. Its presence in the Big Sandy in Kentucky has been established.
All records from outside of this region are to be regarded for the present as doubtful, or positively wrong, as for instance. Grand River, Ontario, and Michigan. The records from Lake Erie given by Sterki (1907a, p. 391) and Ortmann (19096, p. 203) do not refer to this species, but to Pleurobenia obliquum pauperculum, and the same probably is true of Walker’s record (1913, p. 22).
Toward the west and southwest, in the Mississippi-drainage, typical subrotunda is gradually replaced by the form or species F. ebena (Lea), which is, for instance, rather prevalent in the Ouachita River in Arkansas. The interrelation between F. subrotunda and F. ebena should be studied more closely especially in the lower Ohio and its tributaries.
Eusconaia subrotunda kirtlandiana (Lea) (1834).
Quadrula kirtlandiana (Lea) Simpson, 1914, p. 891.
Plate I, figs. 3, 4, 5.
Records from Pennsylvania:
Harn, 1891, p. 137 (western Pennsylvania).
Rhoads, 1899, p. 137 (Beaver River, Wampum, Lawrence Co.).
Ortmann, 1909&, p. 201.
Characters of variety: Shell with the outline of F. subrotunda, but much more compressed, and with less prominent beaks. The diameter is less than fifty percent of the length, falling as low as thirty-three percent. In consequence of the com- pression, the posterior part of the shell, behind the beaks, appears more elevated, almost wing-like. Color of epidermis generally brighter, chiefly so in young shells, which often possess a very light yellowish ground-color, with darker, well- marked growth-rests, and distinct black or dark green rays. In the old shell.
12
MEMOIRS OF THE CARNEGIE MUSEUM.
the color becomes more uniformly brown, and sometimes blackish. All other characters are like those of the main form.
L. H. D. Pr.ct.
Size: Clarksville, Cat. No. 61.3927. .133 mm. 97 mm. 48 mm. .36 (largest at hand). Cochranton, Cat. No. 61.3923 . 128 “ 111 “ 51 “ .40
These sjiecimens are superior in size to the largest F. subrotunda.
The soft parts and glochidia have been described and figured by Ortmann (19116, PI. 89, fig. 1, and 1912, p. 245).
Breeding season: The only gravid female ever found was obtained on August 2, 1908; it had glochidia. Among individuals collected June 25, 1907, and July 10 and 19, 1909, none were gravid, although a good many were obtained. This is rather astonishing. The one gravid specimen was the only one among a large number. Many have been collected later in the season, but no other gravid female has ever turned u]).
Remarks: This form ])asses very gradually into F. subrotunda, and is positively recognized only b,y the compression of the shell. In order to distinguish these two forms I was compelled to introduce an artificial and arbitrary dividing line at the diameter of fifty percent of the length. Of course, this does not correspond to the natural conditions, but it is a wonderful help for the practical separation of the forms.
The other characters are also not reliable, although generally the color and the development of a iiosterior wing help in the identification.
The soft jiarts in kirtlandiana are more frequently of the whitish type, although the orange type is not rare.
In this form it is likewise impossible to distinguish males and females by the shell.
Localities in Pennsylvania represented in the Carnegie Museum:
Ohio River, Industry, Beaver Co.; Coraopolis (S. N. Rhoads) and Neville Lsland, Allegheny Co. (W. F. Graham).
Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).
Mahoning River, Mahoningtown, Coverts, Edinburg, Lawrence Co.
Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsville and Clarksville, Mercer Co. Pymatuning Creek, Pymatuning Township, fiercer Co.
yMlegheny River, Natrona, Allegheny Co.; Godfrey, Johnetta, Kelly, and Templeton, Armstrong Co. French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co. Conneaut Outlet, Conneautlake, Crawford Co.
Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).
Cheat River, Cheat Haven, Fayette Co.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
13
Other localities represented in the Carnegie Museum:
Mahoning River, Ohio (Hartman collection) (Topotype).
Tuscarawas River, Ohio (Holland collection).
West Fork River, Lynch Mines, Harrison Co., West Virginia.
Little Kanawha River, Grantsville, Calhoun Co., West Virginia (W. F. Graham).
Distribution and Ecology in Pennsylvania (See fig. 2) ; The distribution of this form in Pennsylvania clearly indicates that it is the representative of F. suhrotunda in the smaller rivers and creeks, but that it passes in the downstream direction into the latter, and is associated with it in the larger rivers. Its metropolis is in the Beaver system and French Creek. In these it is practically everywhere, and lives in coarser or finer gravel, even in sand, and in more or less rapidly flowing water. It avoids, however, the extreme headwaters, and is not found in the Shenango above Clarksville, and not in French Creek above Cambridge Springs. The records from some smaller tributaries (Pymatuning Creek and Conneaut Outlet) are founded upon single individuals.
In the larger rivers, the Allegheny, Monongahela, and Ohio, it is also present, but its place is gradually taken by the typical F . subrotunda. From Cooks Ferry in Beaver County down the Ohio to Portsmouth, Scioto County, Ohio, the typical form alone is present. Exceptionally large and posteriorly produced specimens may exhibit the dimensions of kirtlandiana , but such are very rare in this section of the river, and generally it is clearly evident that they were typical subrotunda, when young.
General Distribution. Type locality, Mahoning River, Ohio (Lea).
This variety appears to be restricted to the tributaries of the upper Ohio in West Virginia, Ohio, and Pennsylvania. The river which forms its type locality in Ohio also contains it in Pennsylvania, and it occurs also in the other branches of the same river-system (the Shenango and Beaver). I further have ascertained that it is found in French Creek, and the upper Monongahela, its range going down for some distance into the larger rivers. I also found it in the Little Kanawha in West Virginia.
Simpson (1900) quotes it from the “ Ohio, Cumberland and Tennessee River systems, southwest to Arkansas, north to Wisconsin (?), cast through southern Michigan.” I think that this wide range is entirely erroneous. I.iOoking over the literature we find it reported from Ohio (aside from the Mahoning River) from the Ohio itself and some of its tributaries, especially from the Tuscarawas River (Sterki, 1907a). Further it is mentioned from the Grand River in Michigan (Call, 1885, and Walker, 1892 and 1898), and from Waukesha, Waukesha Co.,
14
MEMOIKS OF THE CARNEGIE MUSEUM.
Wisconsin (Call, 1885). This latter locality is doubted by Simpson, and I believe that of all these localities only the Tuscarawas River is reliable.
Outside of the upper Ohio region positive and trustworthy records are absent. Thus, for instance, this shell is missing in the Cincinnati Catalogue of Harper (1896). Sterki’s records from other parts of the state of Ohio are very vague. Call (1896a) does not mention it from anywhere in Indiana, and so fprth. The Michigan record is founded upon Call’s authority, and stands by the side of another one (Wisconsin), which is certainly in error.
Apparently this form has been frequently misunderstood, even by Simpson, and this is the more probable, since corresponding, but not entirely identical, forms are met with elsewhere. A flat form of F. subrotunda is found in Elk River, West Virginia. This I have distinguished as var. leucogona (Ortmann, 19136, p. 89), but in the lower part of this river the connection with typical suhrotunda recurs.
All this tends to show that F. subrotunda has the tendency in the headwaters of the Ohio to develop a flat form, called kirtlandiana, constituting an ecological race of the main species. I wish to call special attention to this; as we shall see further on that similar phenomena present themselves to view in the case of other species.
Eusconaia flava (Rafinesque) (1820).
Quadrula rubiginosa (Lea) Simpson, 1914, p. 872; Quadrula flava (Rafinesque)
Vanatta, 1915, p. 557; Fusconaia flava (Rafinesque) Utterback, 1916,
p. 26.
Plate II, fig. 3.
Records from Pennsylvania :
Clapp, 1895, p. 116 (Allegheny Co.).i^
Rhoads, 1899, p. 137 (Ohio River, Coraopolis, Allegheny Co.).^^
Ortmann, 19096, p. 199.
Characters of the shell: Shell of medium size, but rather heavy. Outline subtrapezoidal. Beaks not very prominent, and not inflated. Beak-sculpture consisting of three to five subconcentric bars, slightly waved, forming an angle upon the posterior ridge, and most distinct there. Often these bars are quite
Marshall (1895, p. 90) quotes this species from the Allegheny in Warren Co., but I consider this an error. Just this instance (and a few others) induce me to question the accuracy of the locality of some of the species recorded from AVarren Co. I have repeatedly collected in this region, but did not find any evidence for the existence of this form.
12 The localities in Allegheny Co. are in the region, where the transition horn, flava into trigona takes place. Some of Rhoads’ specimens should be called flava.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
15
indistinct, even on well-preserved beaks. The fourth and fifth bars are generally marked only by nodular swellings upon the posterior ridge.
Shell rather compressed; diameter less than fifty-five percent of length. Sides flattened, or even with a shallow depression in front of the posterior slope. The latter separated from the sides of the shell by a distinct posterior ridge. Ventral margin from straight to broadly and gently emarginate.
■ Fusconaia flava trigona. • Fusconaia flava,
-j- Fusconaia flava parvula.
Surface without sculpture. Epidermis rather light brown, or of a chestnut or russet hue, rarely showing some green, with fine, indistinct, greenish or brownish rays, and dark growth-rests. When older, the rays disappear, but the epidermis remains rather light, and only in very old shells turns dark brown or even blackish.
Hinge-teeth well-developed; pseudocardinals divergent, rather strong. Inter- dentum present, but not very broad. Beak-cavities not very deep. Nacre white, often suffused with salmon-pink.
L. H. D. Pr.ct.
Size: 1. New Sheffield, Cat. No. 61.3343 95 mm. 71 mm. 44 mm. .46
2. Rosston, Cat. No. 61.2973 65 “ 48 “ 28 “ .43
Soft parts and Glochidia (See Ortmann, 19116, PI. 89, fig. 2, and 1912, p. 241, fig- 4).
On May 24, 1911, in the Little Kanawha River, at Burnsville, Braxton Co., West Virginia, I found among numerous gravid females with normal (red) color
16
MEMOIRS OF THE CARNEGIE MUSEUM.
of the placentae a single one, in which they were pure white. This has remained the only case of this kind. (I have seen over one hundred gravid females.)
Breeding season: Gravid females have been observed on the following dates. May G, 1910; May 9, 1913; May 17, 1910; May 22, 1912; May 24, 1911; May 27, 1908; June 30, 1908; July 3, 1908; July 8, 1907; August 3, 1909; August 10, 1909. In the month of May only eggs were found, at the other dates eggs and glochidia, or only the latter. The species is typically tachytictic, breeding from May to August.
Remarks: This is a rather characteristic shell, but it has been frequently confused with other species, chiefly with Pleurohema obliquum coccineum (Conrad). Nevertheless the peculiar, subtrapezoidal shape, the pale brown or reddish epi- dermis, the flat sides, and distinct posterior ridge, always serve to distinguish it. It is, however, quite variable, and, as we shall see below, three varieties may be distinguished in Pennsylvania, and there are others outside of this state, which have often been regarded as distinct species. In spite of this it has been positively ascertained that these varieties actually intergrade in our region.
Localities in Pennsylvania represented in the Carnegie Museum:
Ohio River, Neville Island, Allegheny Co.
Monongahela River, Elizabeth, Allegheny Co. (D. A. Atkinson & Graf); Charleroi (G. A. Ehrmann) and Millvale, Washington Co. (.J. A. Shafer).
Raccoon Creek, New Sheffield, Beaver Co.
Chartiers Creek, Carnegie, Allegheny Co. (D. A. Atkinson & Graf).
North Fork Tenmile Creek, Amity, Washington Co.; South Fork Tenmile Creek, Waynesburg, Greene Co.
Dunkard Creek, Wiley and hlount Morris, Greene Co.
Allegheny River, Godfrey, Johnetta, and Kelly, Armstrong Co.
Crooked Creek, Rosston and Southbend, Armstrong Co.; Creekside, Indiana Co.
Other localities represented in the Carnegie Musemn:
Lake-drainage:
Genesee River, Chili and Rochester, Monroe Co., New York (R. H. Santens).
Sandusky River, Fremont, Sandusky Co., and Upper Sandusky, AVyandot Co., Ohio (C. Goodrich). Miami-Erie Canal, Lucas Co., Ohio (C. Goodrich).
Swan Creek, Toledo, Lucas Co., Ohio (C. Goodrich).
Maumee River, Defiance, Defiance Co., Ohio (C. Goodrich); Fort AVayne, Allen Co., Indiana (C. Good- rich).
St. Mary’s River, Roclcford, Mercer Co., Ohio (C. Goodrich).
Beaver Creek, AAhlliams Co., Ohio (C. Goodrich).
St. Joseph River, Indiana (B. AValker).
Raisin River, Grape P. O., Monroe Co., and Adrian, Lenawee Co., Michigan (C. Goodrich).
Clinton River, Utica, Alacomb Co., Michigan (B. AA^alker). ^ '
OKTMANN: MONOGEAPH of the naiades of PENNSYLVANIA.
17
Ohio-drainage:
Tuscarawas River, Ohio (Holland collection).
Wolfe Creek, Washington Co., Ohio (W. F. Graham).
Chillicothe, Ross Co., Ohio (Hartman collection).
Ohio Canal, Columbus, Franklin Co., Ohio (Smith collection).
Scioto River, Kenton, Hardin Co., Ohio (C. Goodrich).
Wabash River, New Cory.don, Jay Co., Geneva, Adams Co., and Bluffton, Wells Co., Indiana (C. Good- rich).
Little Kanawha River, Burnsville Braxton Co., West Virginia.
North Fork Hughes River, Cornwallis, Ritchie Co., West Virginia.
Pocatalico River, Raymond City, Putnam Co., West Virginia.
Coal River, Sproul, Kanawha Co., West Virginia.
Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.
Licking River, Farmer, Rowan Co., Kentucky.
Drainage of upper Mississippi and Red River of the North.
Kishwaukee River, Rockford, Winnebago Co., Illinois (P. E. Nordgren).
Sheyenne River, Argusville, Cass Co., North Dakota (S. M. Edwards).
Western and Southwestern Range.
Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).
Wakarusa River, Lawrence, Douglas Co., Kansas (R. L. Moodie donor).
Wea Creek and Bull Creek, Miami Co., Kansas (C. Goodrich donor) (Osage drainage).
Neosho River, Burlington, Coffey Co., Kansas (R. L. Moodie donor).
Terre Noir Creek, Mount Zion, Clark Co., Arkansas (H. E. Wheeler).
Big Deceiper Creek, Gum Springs, Clark Co., Arkansas (H. E. Wheeler).
Chikaskia River, Tonkawa, Kay Co., Oklahoma (F. B. Isely).
Illinois River, Talequah, Cherokee Co., Oklahoma (F. B. Isely).
Blue River, Durant, Bryan Co., Oklahoma (F. B. Isely).
' Note: The specimens from Arkansas and Oklahoma are absolutely indistinguishable from the Pennsylvanian form in shape and anatomy, but have generally a more brilliant, shining, reddish epidermis.
Distribution and Ecology in Pennsylvania (See fig. 3) : In our state F. flava is eminently characteristic of smaller streams, avoiding the larger rivers, although it has been found in the Allegheny, Monongahela, and Ohio. It is most abundant in the southwestern section of the state (Monongahela-drainage), and locally is rather plentiful, as for instance in Raccoon, Tenmile, and Dunkard Creeks. In the Allegheny River it is rather scarce, but is found in considerable numbers in Crooked Creek. Nevertheless elsewhere in the Allegheny-drainage it is absent, which is especially true of French Creek and the uppermost Allegheny, where there is a rich fauna still present, which has been well investigated. It is also absent in the whole Beaver-drainage.
It is hard to say what may have caused this peculiar condition. Yet atten- tion should be called to the fact that the Beaver River and French Creek belong to the Glacial area, and that all creeks in Pennsylvania, in which this species is found, are entirely outside of this area. In the Kiskiminetas, Red Bank, and
18
MEMOIRS OF THE CARNEGIE MUSEUM.
r*
Clarion Rivers the fauna is entirely destroyed, and the few survivals we have in the headwaters of the Kiskiminetas do not include this species.
F. flava prefers fine gravel and sand, and avoids rough bottom and rocks. Its
favorite stations are on bars of fine, firmly packed gravel, just below riffles.
\ ■ .
General Distribution: Type locality, Small tributaries of Kentucky, Salt River, and Green River (Rafinesque).
Outside of Pennsylvania this form has a wide range.- It has been reported from western New York (see below) westward as far as Kansas and southeastern Nebraska. Northward it passes into Canada and the drainage of the Red River of the North (Winniiieg, compare also our specimens from North Dakota). In Michigan, it is all over the southern half of the lower iieninsula (Walker’s map, 1898, PL 1). It probably reached the lake-drainage by several ways, but not through Pennsylvania, since it is missing in the Beaver and upper Allegheny basins.
From the Ohio River southward its range becomes obscure. As I have dis- covered, it is present in the Little and Big Kanawha-drainages in West Virginia, and also in the Big Sandy and Licking Rivers in Kentucky. The type locality is in central Kentucky. Records from Tennessee are missing, except that given by Wilson & Clark (1914) Stones River, Tennessee, tributary to the Cumberland. The localities quoted from Mississippi, Alabama, and Texas are more than doubtful. But, as our material shows, forms representing the species are certainly found as far west as Arkansas and Oklahoma, and although certain authors might call, and have called, these by different names, I am unable to distinguish them from the northern form, except by their more shining epidermis. (See Wheeler, 1918, p. 123.)
It would be interesting to Imow whether outside of Pennsylvania the rule likewise holds good, that this form prefers smaller creeks. This is certainly the case, wherever I have collected it in West Virginia and Kentucky. The localities in Arkansas, and partly also in Oklahoma, are small creeks, while in the larger rivers (Ouachita) Fi flava undata and trigona are found. Call (1900, p. 506) says that rubiginosus (flava) is found in Indiana, in streams both large and small, while Wilson & Clark (1912a, pp. 42, 43) report /am from the headwaters of the Kankakee system in Indiana, while they cite trigona from the lower Kankakee and Iroquois Rivers in Illinois. In the uppermost Wabash and in the Maumee C. Goodrich collected only typical flava, and here undoubtedly is one of the places, where it
“ However, in Elk River in West Virginia this is not perfectly clear. Here is found a form which may be called a dwarfed F. flava trigona. But this is in keeping with the general character of the Elk River fauna, which should be designated as a dwarfed big-river-fauna. It is not the place here to give details of these remarkable conditions,
ORTMANN: monograph of the naiades of PENNSYLVANIA.
19
crossed into the lake-drainage. There is no doubt that jiava has often been con- founded with trigona (or even undata), and that it actually intergrades through trigona into undata.
Fusconaia flava trigona (Lea) (1831).
Quadrula trigona (Lea) Simpson, 1900, p. 787; Quadrula undata (Barnes) (Ohio-
form) Walker, 19106, p. 22; Quadrula undata (Barnes) Simpson, 1914,
p. 880 (pro parte).
Plate II, fig. 1.
Records from Pennsylvania:
Stupakoff, 1894, p. 135 (Allegheny Co.).
Rhoads, 1899, p. 137 (Ohio River, Coraopolis, Allegheny Co.).^^
Characters of variety: This is a Fusconaia flava which has a more swollen shell, chiefly anteriorly, with a diameter of fifty-five percent of the length or more. In consequence of this the sides of the shell are generally more concave, forming a gentle radial depression in front of the posterior ridge. In other respects there are hardly any differences from the normal form.
L. H. D. Pr.ct.
Size: 1. Neville Island, Cat. No. 61.18376 67 mm. 52 mm. 38 mm. .57
2. do. Cat. No. 61.16336 50 “ 41 “ 31.5 “ .63
According to Lea’s figure 59 “ 49 “ 38 “ .64
The soft parts have never been observed in Pennsylvania. But specimens referable to this form have been found with the soft parts in Elk River, West Virginia, and gravid females were found there on July 8, 1911, with glochidia. The anatomy is absolutely identical with that of F . Hava, as are also the glochidia: L. 0.15 mm., H. 0.16 mm. A form indistinguishable from this was collected by H. E. Wheeler in Saline River, Arkansas (July 13, 1911), and the anatomy and glochidia of this form were the same.
According to the above dates, the end of the breeding season of this form falls in July.
Remarks: In this case also I have been compelled to draw an artificial dividing- line, at the diameter of fifty-five percent, between two forms, while in nature a gradual transition exists. This is justified by the same practical considerations as in the case of Fusconaia subrotunda and kirtlandiana. Walker specifically unites the present form with undata and he is undoubtedly right. Yet I think we should recognize trigona as a distinct variety, with less developed beaks; while F. flava undata has much elevated and often incurved beaks. The range of the two forms
I have (19096, p. 183 and 187) questioned the correctness of this record, since Rhoads’ specimens are too young. But it should stand,
20
MEMOIRS OF THE CARNEGIE MUSEUM.
also seems to be somewhat different, F, (lava undata (Barnes), belonging to the larger rivers of the central basin, being absent in Pennsylvania. In the middle West (Illinois) these two forms seem, however, to overlap, and in the Southwest also both seem to be present, and, according to what Walker says, intergrades are present. The real F. flava undata is, in consequence of the higher beaks, more subtrigonal (not subtrapezoidal) in outline, and in addition has the tendency in the epidermis to become greenish rather than brownish.
From the measurements given above we see that some of our specimens agree rather closely with Lea’s figure of U. trigonus. Others reveal transitions in the direction of F. flava. Specimens from Charleroi have been determined by Simpson as intergrades between trigona and ruhiginosa (= flava), and this is entirely correct. We have here (or rather had) in the Ohio below Pittsburgh, and in the Monongahela above, a region, where flava gradually passes into trigona. Further upstream, chiefly in the Monongahela system, only more or less typical F. flava arc found. The only way to bring order out of the chaos is to draw an artificial line, as I have done.
Localities in Pennsylvania represented in the Carnegie Museum:
Ohio River, Neville Island, Allegheny Co.
Monongahela River, Westmoreland Co. (G. A. Ehrmann); and Charleroi, Washington Co. (G. A. Ehr- mann).
Other localities represented in the Carnegie Museuyn:
Elk River, Sutton and Gassaway, Braxton Co., and Shelton, Clay Co., West Virginia.*®
West Fork White River, Riverside, Greene Co., Indiana (J. D. Haseman).
Wabash River, New Harmony, Posey Co., Indiana (A. A. Hinkley).
(A form indistinguishable from this has been received from various localities in the Southwest; but they cannot be distinguished from U. chuni Lea.)
Marais des Cygnes River, Rich Hill, Bates Co., Missouri (W. I. Utterback).
Cache River, Nemo, Craighead Co., and Sedgwick, Lawrence Co., Arkansas (H. E. Wheeler).*’
White River, Cotter, Baxter Co., Arkansas (A. A. Hidkley).
Saline River, Benton, Saline Co., Arkansas (H. E. Wheeler).*®
Kiamichi River, Tuskahoma, Pushmataha Co., Oklahoma (F. B. Isely).
Sabine River, De Soto Parish, Louisiana (L. S. Frierson).
Distribution and Ecology in Pennsylvania (See fig. 3) : F. flava trigona is found in Pennsylvania only in the Ohio and lower Monongahela Rivers, and reaches here
*® Smaller than the Pennsjdvanian form, but agreeing in all other respects. A few specimens from Gassaway would fall under typical flava, but the diameter remains above fifty percent.
*’ In part cotypes of F. selecta Wheeler (Nautilus, XXVIII, 1914, p. 76, PI. 4). I cannot separate them from this form.
*® In some of these, the diameter falls below fifty-five percent, but remains above fifty percent.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
21
the uppermost limit of its distribution in the Ohio system. It is remarkable that I have not found this form or any representative of it in the Ohio in Beaver County. Its absence in the lower Allegheny may be accounted for by the general destruction of the molluscan fauna in these waters. Aloreover F. flava trigona probably is extinct in the state at the present time. The specimens collected by myself at Neville Island were all dead, and there are no more shells at this locality, nor at Rhoads’ locality, Coraopolis. Ehrmann’s collections in the Monongahela were made before 1898, and most of his shells were found dead.
General Distribution. Type locality: Ohio River, Cincinnati and Louisville (Lea).
The distribution of F. flava trigona is very unsatisfactorily known. Walker reports it from the Ohio, and it extends westward to the Mississippi at Davenport, Iowa, and to the Wisconsin River, Sauk Co., Wisconsin (Walker).
The Carnegie Museum possesses a number of specimens from the Kishwaukee River, Rockford, Winnebago Co., Illinois (P. E. Nordgren), which in general correspond with F. flava; but one among them, which is considerably swollen, might very well be considered to belong to the variety trigona. These shells are rather large, with blackish epidermis. The other localities mentioned above, are rather isolated (in West Virginia, Indiana, Arkansas, Oklahoma, Louisiana), but they tend to show that under certain conditions, the form flava undata passes, in localities remote from each other, into a form with less elevated beaks, which answers to the description of U. trigonus of Lea. What these conditions are, remains to be seen. Possibly trigona is the form of medium-sized rivers with strong currents.
It is remarkable that I did not see a trace of this form on my collecting trips down the Ohio between Pittsburgh and Cincinnati. The shell should be expected in this region, but careful examination pf the shell-heaps of the clam-diggers and my own collecting did not bring to light a single specimen. Probably this form selects particular stations in the river, but of what character these are, is as yet unknown. At Neville Island in Pennsylvania, I found the dead shells in and above riffles in a small branch of the Ohio, immediately below a rather long, quiet pool.
Fusconaia flava parvula Grier (1918).
Unio rubiginosus Norris, 1902, p. 119 (Winona Lake); Quadrula rubiginosa Ortmann, 19096, p. 203 (Lake Erie); Quadrula undata (pars) Walker, 19106; Quadrula undata {pars) Simpson, 1914, p. 880; Fusconaia flava parvula Grier, 1918, p. 11.
22
MEMOIRS OF THE CARNEGIE MUSEUM.
Plate II, fig. 2.
Previously reported as a distinct form, only by Grier (1918) but referred to by Ortmann, 19096, p. 203, as Quadrula ruhiginosa.
Characters of variety: Much smaller than F. flava and F. flava trigona, generally less than half their size and bulk. The shape of this form is much like that of the var. trigona (subtrapezoidal) , although sometimes there are found more triangular specimens, which incline towards the western F . flava undata. With regard to the swelling of the valves, there is great variety, some specimens being almost as flat as the normal F. flava, but on the average, the swelling, and also the development of the beaks, is more like that of F. flava trigona, with the diameter generally over fifty percent of the length. Color of epidermis, when young, rather light, yellowish brown (in Lake Erie), or greenish brown (in Winona Lake), with very distinct and regular, dark growth-rests, and fine, indistinct greenish or brownish rays. Old shells become darker, chestnut-brown or greenish black (the latter is the case in Winona Lake). Old specimens are sometimes unusually drawn out at the lower posterior end, and thus become oblique.
|
L. |
H. |
D. |
Pr.ct. |
||||
|
1. |
Erie, |
Cat. No. |
61.3886 |
59 mm. |
49 mm. |
31 mm. |
.53 |
|
2. |
do. |
Cat. No. |
61.4371 |
,57 “ |
49 “ |
33 “ |
.58 |
|
3. |
do. |
Cat. No. |
61.4513 (T3'pe set) |
52 “ |
41 “ |
27 “ |
.52 |
|
4. |
do. |
Cat. No. |
61.4370 |
41 “ |
35 “ |
23 “ |
.56 |
The first two shells are very old, the largest at hand, and they are much drawn out posteriorly; the third and fourth are good average specimens.
Soft parts: They have been alluded to by Ortmann (1912, p. 241), under F. ufidata. A gravid female with young glochidia was subsequently secured from Cedar Point, Ohio, and, as far as could be made out, the glochidia agree in shape and size with those of F. flava.
Breeding season: Gravid females were found in Presque Isle Bay on July 8, 9, and 12, 1910. They did not have glochidia. The specimen from Cedar Point with young glochidia was collected July 24, 1911.
Remarks: This is the Lake Erie .form of F. flava, and a similar form is found in Winona Lake in Indiana. The latter agrees in size, and also in general shape, although the tendency toward the triangular shape of F. flava undata is more pronounced. It is, however, more greenish brown in color, while specimens from Lake Erie are more yellowish or rusty brown. One specimen from Winona Lake is as flat as F. flava.
The variety from Lake Erie is quite distinct, characterized chiefly by its small, dwarfed size, and by having more distinct and regular growth-lines, a
ORTMANN: monograph op the naiades of PENNSYLVANIA.
23
feature noticeable in almost all species found in Lake Erie. In other respects this variety is rather indifferent; it inclines most toward the var. trigona, but there is a tendency in some individuals to become more triangular (with higher beaks) ^ like F. flava undata. With regard to obesity, it is very variable, and some specimens are as flat as F. flava; in fact, young specimens are very often found, which are indistinguishable from young F. flava.
Localities represented in the Carnegie Museum:
Lake Erie, Presque Isle Bay, Erie, Erie Co., Pa.; and also in Horseshoe Pond on Presque Isle.
Lake Erie, Crystal Beach, Welland Co., Ontario, Canada (F. Behrle); Port Rowan, Norfolk Co., On- tario, Canada (C. Goodrich); Sandusky Bay, Cedar Point, Erie Co., Ohio (0. E. Jennings); La
Plaisance Bay, Monroe Co., Michigan (C. Goodrich).
Winona Lake, Kosciusko Co., Indiana (E. B. Williamson) .1®
Distribution and Ecology (See fig. 3) : Type locality, Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania. Type set: Carnegie Museum, Cat. No. 61.4513.
Aside from the range indicated by the above localities, no details are known. Walker (19105, p. 22) mentions that Quadrula undata has invaded the St. Lawrence system, and that it is found in the lake-drainage in Wisconsin, Illinois, and southern Michigan, and that it goes eastwards as far as Buffalo, New York, and Port Dover, Ontario (in Lake Erie); and in 1913, Walker cites both rubiginosa and undata from Lake Erie, but the form from Lake Erie has not been discussed in detail. We do not know whether the var. parvula is restricted to the lake, or also found elsewhere. Marshall (1895, pp. 89 and 93) calls the Lake Erie form from New York both rubiginosa and trigona.
As far as my material goes (I have seen over fifty specimens from Lake Erie) F. flava parvula is a well-marked local form, easily distinguished from the other phases of the species. However, it would not be astonishing if it should pass else- where into the other forms, in fact, the shells from Winona Lake represent to a degree transitions both toward F. flava and F. flava undata.
Possibly the dwarf form of Quadrula rubiginosa from Tippecanoe and Kuntz Lakes in northern Indiana, mentioned by Wilson & Clark (1912a, p. 43), difficult to distinguish from trigona, belongs here.
The mutual connection of the three forms described above has hitherto been misunderstood, and some authors have been quite emphatic in the assertion that rubiginosa ( = flava) should not be united specifically with undata or trigona (Call, 1895; Walker, 19106). But upon the basis of my own studies of the conditions in western Pennsylvania, I am as emphatic in maintaining that flava is only the
These were received as “ coccinea,” but undoubtedly correspond to “ rubiginosa ” of Norris’ list (1902) from Eagle (= Winona) Lake.
24
MEMOIRS OF THE CARNEGIE MUSEUM.
creek-form of the river-form flava trigona. As I have pointed out under jiava trigona, the latter has (or had) in the Ohio and Monongahela near Pittsburgh a distinct tendency to liecome more flattened. There are all intergrades in this respect before me, and it is absolutely impossible to draw a natural line between the swollen trigona-iorm, and the flat yZaya-form. Farther up in the Monongahela and Allegheny, and in their tributaries, and also in the smaller tributaries of the Ohio jiroper, the pure and typical F. iiava is exclusively found.
Conditions like these force us to unite these forms specifically. But never- theless trigona is to be regarded as a distinct variety, constituting a geographical, dr rather ecological, race of F . flava, which has its definite habitat in larger streams. There are indications that similar conditions prevail outside of Pennsylvania.
Attention should be called to the singular iiarallelism of these forms with those of the Fusconaia subrotunda-gvo\\\> . In the latter, we have seen that there is a greatly swollen form, with highly elevated and incurved beaks (F. ebena) living in the largest, deepest rivers, with muddy bottoms, and therefore in those regions, which are nearest to the center of the interior basin. In the great rivers with sandy and gravelly bottoms and somewhat s,tronger current, that is to say in the upiier Ohio, this is replaced by a form, which, although more or less swollen, has less elevated lieaks (F. suhrotunda) . This in turn in the smaller streams gives way to a flat form (F. kirtlandiana) . The same is true apparently in the F. flava-groiiY). We observe a very swollen form with high beaks having the Mississipjii as the center of its range (F. undata) ; a swollen form with less elevated beaks in the upjier Ohio (F. trigona) ; and a flat form in the smaller creeks (F. iiava) . To these is added, in this case, a dwarfed lake-form in Lake Erie (F. parvula).
The boundaries of these corresponding forms of the two series do not coincide, inasmuch a F. subrotu7ida goes farther up in the rivers than does F. flava trigona.
It is well to keep this peculiar phenomenon in mind, for later on we shall liecome acquainted with other instances of the same kind.
Further, attention should be called to the peculiar fact that, while the form flava is entirely missing in northwestern Pennsylvania, and is represented in Lake Erie by the form parvula, it turns up again in western New York. The Carnegie Museum has specimens from the environs of Rochester, New York, and it had been previously reported from the region of Buffalo and the Erie Canal, from the Genesee River and the Mohawk, crossing over to the Atlantic slope (See Call, 1878, and 1885; Dewey, 1856; Marshall, 1895; Baker, 18985). According to Call, it has migrated along this route in recent times. Since this range has no connection with the rest of the range of F. flava, yve must assume that it came
ORTMANN: monograph op the naiades of PENNSYLVANIA.
25
into this region from Lake Erie near Buffalo, and that it is the form 'parvula which originally entered the tributaries and canal, but which under the changed environ- ment, again assumed the shape and the characters of the form of the creeks, called Hava. This is a very interesting phenomenon, and appears to indicate the very important influence of environment upon the shape of the external characters of the shell. If this were the only instance of the kind, we might regard it with suspicion, but we shall become acquainted with other facts of a similar nature, and in order not to loose sight of this instance, it is emphasized here.
Finally attention should be directed to the great tendency to develop local forms within the /Zawa-groiip. Although we have distinguished only four main forms {undata, trigona, flava, parvula) some of these include several local types. For instance, the flava of the Allegheny River and Crooked Creek in Pennsylvania is distinguished by small size, while in the creeks of southwestern Pennsylvania a much larger form prevails. In the Little Kanawha River in West Virginia is a very peculiar race of flava, distinguished by a shape, which is considerably drawn out at the lower posterior end. A peculiar small race of trigona is found in Elk River, West Virginia, and the forms from Arkansas also have certain charac- teristics of their own. These conditions are very interesting, but can be studied only with the help of larger series from the different localities.
Genus Amblema Rafinesque (1820).
Crenodonta Ortmann, 1912, p. 245; Crenodonta (Schlueter) (section of Quadrula) Simpson, 1914, p. 813; Amblema Frierson, 1914rt, p. 7; Amblema Utter- back, 1916, p. 31.
Type Amblema costata Rafinesque.
Only one species, A. plicata, is found in Pennsylvania, within which, however, two well-marked races may be distinguished.
Key to the forms of A. plicata.
«!. Shell smaller, generally more swollen, less elevated posteriorly. Sculpture less developed, chiefly
so upon the posterior slope A. plicata.
a 2. Shell larger, generally more compressed, and more elevated posteriorly. Sculpture better developed, chiefly so upon the posterior slope A. plicata costata.
Amblema plicata (Say) (1817).^^’
Quadrula plicata hippopcea (Lea) Simpson, 1914, p. 816; Amblema plicata (Say) Utterback, 1916, p. 33.
“Not 1816, as given by Simpson. (See Binney, Bibliography, 2, Smithsonian Miscell. Coll., 9, 1869, p. 277.)
23
MEMOIRS OF THE CARNEGIE MUSEUM.
Plate II, figs. 4, 5, 6.
Records from Pennsylvania :
Reported previously from Lake Erie, but not from the Pennsylvanian shores, except by Ortmann (19096, p. 203) as undulata hippopcea.
Characters of the Shell: Shell of medium size, rather heavy. Outline suboval to subtrapezoidal, generally slightly longer than high, somewhat oblique, rounded before, subtruncate behind. Beaks moderately prominent. Beak-sculpture con- sisting of three to five concentric ridges, slightly angular and nodulous behind; only one to three are distinct, the others indistinct, often indicated only by the nodules. The beak-sculpture is not continued upon the disk. Shell moderately swollen, with indistinct, rounded posterior ridge. Surface sculptured by trans- verse, oblique folds or undulations, which run parallel to each other in the direc- tion of the lower posterior end. These folds are absent in very young shells, and begin in older individuals at a certain distance from the beaks, and at, or a little in front of, the middle of the disk, leaving the anterior part of the shell free. These folds are very variable, sometimes hardly indicated, in other cases they are rather strong. In large shells, there are as many as four or five of them. Posterior slope comparatively narrow, since the upper posterior margin is only little, or not at all elevated, so that the posterior wing is only slightly developed. In younger shells, however, it is generally well-developed. The posterior slope is often entirely smooth, but sometimes there are indications of radiating folds or ribs, smaller than those upon the middle of the disk, and occasionally they may become rather distinct and strong. In the latter case they have no distinct relation to the large folds, but seem to diverge from the uppermost of the latter at an angle. Epi- dermis in young specimens yellowish or greenish brown, rather light, without rays. Growth-lines dark. In old shells the epidermis becomes darker, reddish or chestnut-brown to blackish, and the growth-rests, which are rather regular, become less marked in color.
Hinge-teeth strong, ragged. Pseudocardinals divergent. Interdentum mod- erately developed. Lateral teeth strong and rather long. Beak-cavity of medium depth. Dorsal muscle-scars on the hinge-plate. Nacre white, posteriorly often very beautifully iridescent with purplish and pinkish reflections. No difference whatever between the male and female shell.
|
L. |
H. |
D. |
Pr.ct. |
||||
|
Size: I. |
Erie, |
Cat. No. 61.4520 (largest, 9 ) . • • • |
98 mm. |
71 mm. |
40 mm. |
.41 |
|
|
2. |
do. |
Cat. No. 61.4520 |
(c^) |
68 “ |
54 “ |
30 “ |
.44 |
|
3. |
do. |
Cat. No. 61.4516 |
(cf) .... |
46 “ |
36 “ |
20 “ |
.43 |
*
ORTMANN: monograph of the naiades of PENNSYLVANIA.
27
Soft parts (See Ortmann, 1912, p. 246). Glochidia unknown.
Breeding season: Of this form, I have found only two gravid females, with eggs, on July 8, 1910. The eggs were white.
Remarks: The nomenclature of this species and its forms has been erroneous hitherto, and the original U. plicatus of Say has been entirely misunderstood by practically all authors, except Utterback (1916). Indeed, from Say’s description alone it is impossible to decide what form of the plicata-group he had before him, but the locality he gives, Lake Erie, settles the case. There is no other form of the plicata-group in the lake,^^ except this, and consequently the common form from Lake Erie should bear this name, and not that of hippopcea given by Lea.
A. plicata of Lake Erie is not the normal type of the species, but is a dwarfed form of a species of the Ohio-drainage {costata). Thus nomenclature reverses the natural conditions, making out of a local race the typical form of the species, but this cannot be helped. The differences of A. plicata from the Ohio-form costata are very slight, in fact, the only reliable distinction is the size, but there are a few others, which may be more or less depended upon. The typical plicata is generally more oval, and the posterior part (wing) is not so elevated. Furthermore the growth-lines are more regular and closer together, the shell is slightly more swollen on the average, and the color lighter. But in young specimens all these characters are more or less obscured, and it is practically impossible to distinguish these two forms in the juvenile stage. I have young costata which may be matched with young plicata, without showing the slightest difference.
Localities represented in the Carnegie Museum:
Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania.
Lake Erie, Cedar Point and Sandusky Bay, Erie Co., Ohio (O. E. Jennings and Chas. Brookover).
Lake Erie, La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).
Distribution and Ecology (See fig. 4). Type locality: Lake Erie (Say).
This species is known in Pennsylvania only from Lake Erie. In Presque Isle Bay it is rather abundant, and at certain places sometimes great numbers of young shells are found in one to two feet of water. The larger specimens are generally found at a depth of three to four feet, and with the clam-dredge I obtained some at five feet, and others were brought up by the “ sand-sucker ” from a depth of ten to fifteen feet. They are found generally in fine sand and gravel, but I also found a few on the southern shore of the bay (mainland), where the bottom consists of coarse shingle.
1 have received, from L. S. Frierson, a true A. plicata costata from Maumee Bay at the mouth of the Ottawa River, near Toledo, Lucas Co., Ohio. This however, is a dead shell, and may have been washed into the lake from the Ottawa or Maumee Rivers, in both of which A. plicata costata is present.
28
MEMOIRS OF THE CARNEGIE MUSEUM.
Outside of Pennsylvania this species has been recorded only from those states which border on Lake Erie. It is known from Buffalo, New York (Marshall, 1895, p. 50), but said to be identical with U. undulatus {= A. costata). It is given
• Amblema flicata.
■ Amhlema plicata costata.
from the Ohio shores of the lake by Sterki (1907a), and from the shores in south- eastern Michigan (Walker, 1898), and from Lake Erie in general (Walker, 1913).
The record from Lake Winnipeg, Canada (Hanham, 1899) should be questioned for the present.^^
Amblema plicata costata (Rafinesque) (1820).
Quadrula undulata (Barnes) Simpson, 1914, p. 819 Quadnda costata (Rafin- esque) Vanatta, 1915, p. 556; Aynblema 'plicata costata (Rafinesque) Utterback, 1916, p. 33.
Plate II, fig. 7; Plate III, figs. 1, 2, 3.
Records from Pennsylvania:
Clapp, 1895 (Allegheny Co.).
Marshall, 1895 (Allegheny River, Warren Co.).-''
Rhoads, 1899 (Ohio River, Beaver Co., and Beaver River, Wampum, Lawrence Co.) (as plicatus). Ortmann, 19095, p. 198.
22 Simpson (1900, p. 769, footnote 3) mentions a peculiar form of “ undulata ” from Lake Winni- peg. Possibly this is the same as that referred to by Hanham, and might be a form parallel to plicata.
2^ Simpson makes a singular mistake in quoting, among the synonyms, Unio undulatus Sowerby, 1868, PI. 76, fig. 399, which probably is Alasmidonta undulata (Say).
2-* I did not find this form so far up in the Allegheny, but only as high up as Venango Co., where it is very rare. But it may have e.xtended farther up in the past.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
29
Characters of variety: This form differs from plieata of Lake Erie by being considerably larger, attaining a huge size; in fact it is one of our largest and heaviest shells. The shell is generally rather flat, with the diameter less than forty percent of the length, and the posterior wing is well-developed, so that the upper margin appears elevated posteriorly. The beaks are very slightly prominent. The surface sculpture is similar to that of plieata. In large specimens there are upon the sides four or flve transverse bars, but they are, as a rule, more distinct and broader than in A. plieata. The radiating ribs upon the posterior slope and the wing are more frequently and more distinctly developed. Color of epidermis generally darker, although young shells are often as light as plieata. The growth-lines are less regular, and old shells become uniformly black. Nacre white, often beauti- fully iridescent posteriorly. No difference between the male and the female in the shell.
L. H. D. Pr.ct.
Size: 1. Jamestown, Cat. No. 61.3831 161 mm. 123 mm. 68 mm. .42
2. Mahoningtown, Cat. No. 61.3830 99 “ 78 “ 38 “ .38
No. 1 is the largest specimen on hand.
Soft parts and Glochidia (See Ortmann, 1912, p. 246).
Breeding season: Typically tachytictic, breeding from May to July. I found gravid females on the following dates: May 13, 1911; May 17, 1910; May 23, 1911; May 23, 1912; May 23, 1914; May 24, 1911; June 19, 1909; July 3, 1908; July 8, 1909; July 10, 1909; July 10, 1911. Glochidia were seen only in July, and a discharging specimen was observed on July 10, 1911. Wilson & Clark (1912a) report gravid specimens as late as July 28.
Remarks: Amhlema plieata costata undoubtedly is the parent form, from which A. plieata is derived as a depauperate form in Lake Erie. The former is rather variajble in some of its characters, but it is always much larger, generally more compressed, and the posterior wing is more elevated. The sculpture varies a good deal. I have specimens in which the shell is practically smooth, without any folds (Compare the flgures of Baker, 1898a, PI. 12, fig. 1; Smith, 1899, PI. 82, and our figs. 2 and 3 on plate III). Sometimes the folds are peculiarly developed, showing a tendency to be divided into nodes. I have even a specimen in which these nodes form irregular vertical ribs, running toward the lower margin of the shell, so that, together with the ribs of the posterior slope, a system of low bars is indicated, which diverge from the posterior ridge of the shell in the direction of the lower and posterior margins.
The specimens from Pennsylvania are all to be regarded as true costata, and I have not seen any forms which incline toward the western and southwestern
30
MEMOIRS OF THE CARNEGIE MUSEUM.
representatives; nevertheless there is a tendency in this form to become more swollen in the larger rivers (See PL III, figs. 2, 3).
Localities in Pennsylvania represented in the Carnegie Museum:
Ohio River, Shippingport, Cooks Ferry, and Industry, Beaver Co.
Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).
Mahoning River, Mahoningtown, Coverts, Edinburg, and Hillsville, Lawrence Co.
Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsville, Clarksville, Shenango, and Jamestown, Marcer Co.; Linesville, Crawford Co.
Connoquenessing Creek, Ellwood City, Lawrence Co. (G. H. Clapp, H. H. Smith, & G. L. Simpson, Jr.);
Zelienople, Harmony, Butler Co.
Slippery-rock Creek, Wurtemberg, Lawrence Co.
Pymatuning Creek, Pymatuning Township, Mercer Co.
Padan Creek, Linesville, Crawford Co. (0. E. Jennings).
Allegheny River, Aladdin, Godfrey, Johnetta, Kelly, and Mosgrove, Armstrong Co.; Walnut Bend, ^ enango Co.
Crooked Creek, Rosston, Armstrong Co.
French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co. Conneaut Outlet, Conneautlake, Crawford Co.
Leboeuf Creek, Waterford, Erie Co.
Dunkard Creek, Dunkard, Mount Morris, Greene Co.
Cheat River, Cheat Haven, Fayette Co.
Locality in Pennsylvania represented in the Philadelphia Academy of Natural Sciences:
Ohio River, Beaver, Beaver Co. (S. N. Rhoads).
Other localities represented in the Carnegie Museum:
Lake-drainage:
Tonawanda Creek, Erie Co., New York (Smith collection).
Sandusky River, Upper Sandusky, Wyandot Co., Ohio (C. Goodrich).
IMaumee River, Roche de Boeuf Rapids and Otsego Rapids, Wood Co., Ohio (C. Goodrich).
Swan Creek, Toledo, Lucas Co., Ohio (C. Goodrich).
Silver and Beaver Creeks, Williams Co., Ohio (C. Goodrich).
St. Mary’s River, Rockford, Mercer Co., Ohio (C. Goodrich).
Ottawa River, Lucas Co., Ohio (C. Goodrich)..
Lake Erie, Maumee Bay, at mouth of Ottawa River, Lucas Co., Ohio (L. S. Frierson donor).^^
Raisin River, Grape P.O., Monroe Co., and Adrian, Lenawee Co., Michigan (C. Goodrich).
Rouge River, Wayne Co., Michigan (C. Goodrich).
Gratiot Co., Michigan (C. Goodrich) (Saginaw drainage).
Ohio-drainage:
Ohio River, Toronto, Jefferson Co., Ohio; St. Mary’s, Pleasants Co., and Parkersburg, Wood Co., West Virginia; Portland, Meigs Co., Ohio.
Conotton Creek, New Hagerstown, Carroll Co., Ohio.
Tuscarawas River, Ohio (Holland collection).
A dead shell, probably washed into the lake from the river.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
31
Wolfe Creek, Washington Co., Ohio (W. J. Graham).
Ohio Canal, Columbus, Franklin Co., Ohio (Smith collection).
Scioto River, Kenton, Hardin Co., Ohio (C. Goodrich).
Big Beaver Creek, Mercer Co., Ohio (C. Goodrich).
Wabash River, New Corydon, Jay Co., Geneva, Adams Co., and Bluffton, Wells Co., Indiana (C. Good- rich).
West Fork River, Lynch Mines, Harrison Co.; Milford, Harrison Co. (W. J. Graham); Lightburn and Weston, Lewis Co., West Virginia.
Little Kanawha River, Grantsville, Calhoun Co. (W. J. Graham) and Burnsville, Braxton Co., West Virginia.
North Fork Hughes River, Harrisville (W. J. Graham), and Cornwallis, Ritchie Co., West Virginia.
Elk River, Shelton, Clay Co., and Gassaway and Sutton, Braxton Co., West Virginia.^®
Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.
Licking River, Farmer, Rowan Co., Kentucky.
Tennessee-drainage :
French Broad River and Boyd Creek, at Boyd Creek, Sevier Co., Tennessee.
Nolichucky River, Chunns Shoals, Hamblen Co., Tennessee.
Holston River, Hodges, Jefferson Co.; Turley Mill and Noeton, Grainger Co.; Austin Mill, Hawkins Co., Tennessee.
South Fork Holston River, Pactolus, Sullivan Co., Tennessee.
North Fork Holston River, Rotherwmod, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia.
Clinch River, Solway, Knox Co.; Clinton and Offutt, Anderson Co.; Black Fox Ford, Union Co.; Clinch River Station, Claiborne Co.; Oakman, Grainger Co., Tennessee; Speers Ferry and Clinchport, Scott Co., Virginia; St. Paul, Wise Co., Virginia; Fink and Cleveland, Russell Co., Virginia. Emory River, Harriman, Roane Co., Tennessee.
Powell River, Combs, Claiborne Co., Tennessee.
From upper Mississippi northwards and westwards:
Little Muddy Creek, DuBois, Washington Co., Illinois (A. A. Hinkley).
Sheyenne River, Argusville, Cass Co., North Dakota (S. M. Edwards) (drainage of Red River of the North).
Meramec River, Meramec Higlilands, St. Louis Co., Missouri (N. M. Grier).
James River, Galena, Stone Co., Missouri (A. A. Hinkley).
Terre Noir Creek, Mount Zion, Clark Co., Arkansas (H. E. Wheeler).
All the above specimens are to be considered as typical A. plicata costata. Other sets from Indiana, Illinois, northern Alabama (Tennessee-drainage), Kansas, Arkansas, and Oklahoma, show peculiar features, leading to the western and southern representatives of this form. I cannot go into detail here, yet it should be mentioned that I have from the Alabama drainage one set, which cannot be distinguished from costata. It is from Valley Creek, Toadvine, Jefferson Co., Alabama (H. H. Smith).
Distribution and Ecology in Pennsylvania (See fig. 4) : This form is not rare in our state. It is found in the larger streams as well as in their tributaries, and in general is most abundant in some of the latter. In the Ohio below Pittsburgh it is not very plentiful; in the Monongahela proper it has not been found, and it is decidedly rare in the Allegheny. But it turns up again in some of the tributaries
In the Kanawha drainage, I have seen a dead shell in Coal River, Sproul, Kanawha Co., W. Va.
32
MEMOIRS OF THE CARNEGIE MUSEUM.
of the Monongahela, and is abundant in its headwaters in West Virginia. In the Beaver-drainage it is nearly everywhere, being alisent only in the smaller creeks. The same is true of French Creek.
It is generally found in the more or less coarse gravel of our streams, often in the Allegheny and Ohio among very heavy gravel and in strong currents, but it is by no means averse to fine gravel and sand (chiefly so when young). It avoids mud, as most of our Naiades do. The shell attains considerable size, and there is no relation between the size of the stream and that of the shell, very large individuals having been often found in small creeks.
General Distribution: Type locality, Ohio River (Rafinesque) ; according to Vanatta small creeks in Kentucky.
The range outside of Pennsylvania is very extended, and comprises practically the whole of the Ohio-drainage, including the Tennessee-Cumberland system, and westward, the Mississippi- and Missouri-drainages to Minnesota, Iowa, Ne- baska, Kansas, and Oklahoma. Northwards it crosses into the drainage of the Great Lakes at several points, and has been reported as occurring as far northeast as Ottawa, Canada (Simpson, 1893, p. 592). It is found in western New York in Erie, Niagara, Alonroe, and Onondaga Cos. (Marshall, 1895) in tributaries of the St. Lawrence system and along the route of the Erie canal. This group of localities, confirmed also by our specimens from Tonowanda Creek, is important (see below). It is not found in Lake Erie proper, being there replaced by A. plicata.
In Ohio it is present in both drainages (Sterki, 1907a), and has been repeatedly reported from streams running to Lake Erie; from the Cayuga, Rock, and Sandusky Rivers (Dali & Simpson, 1895), and also from the Maumee River (Carnegie Mu- seum). This establishes one route of migration from the Ohio to the lake. It is furthermore found in southern Michigan (Walker, 1898). It also crosses over in the north into the Red River of the North and the Lake Winnipeg-drainages.
Towards the south and southwest, the boundaries of the range of the true A.
¥
plicata costata are poorly known. It certainly exists in tributaries of the Cumber- land in Tennessee (Wilson & Clark, 1914), and in the headwaters of the Tennessee in eastern Tennessee and Virginia (Carnegie Museum). But farther in this direction it is replaced by the southern form {perplicata) . In Arkansas and Okla- homa forms intergrading with the latter are found.
From the middle Ohio (region of Cincinnati) downward, and in the Mississippi River, its place is largely taken by the closely allied Amhlema peruviana (Lamarck) This is the Quadrula ylicata of Simpson (1914, p. 814).
ORTMANN; monograph of the naiades of PENNSYLVANIA. 33
but particulars about the distribution of this form are scarce. As far as can be judged, A. peruviana seems to prefer the largest rivers, and probably muddy bottoms. In the Ohio from the Pennsylvania state line down to Portsmouth, Ohio, I have never seen a true peruviana.
In the distribution of the forms of the A. plicata-growp two facts should be especially emphasized.
1. We have here a group of at least three forms: a dwarfed form from Lake Erie (A. plicata) ; a flat form found in the smaller rivers and headwaters {A. plicata costata) ; and a swollen form with high beaks in the largest rivers {A . peruviana) . Only the first two are found in Pennsylvania, but the third turns up in the Ohio in the neighborhood of Cincinnati. These conditions correspond in a degree to what we have observed in two cases in the genus Fusconaia (See above, pp. 14 and 24) .
2. In western New York, we have the typical form from small rivers, (costata) tributaries of the St. Lawrence system, apparently entirely isolated from the rest of the range, for this form is positively absent in the uppermost Allegheny- drainage. The form found in Lake Erie is not this, but A. plicata. Thus it seems that western New York has been colonized from the lake, the lake-form (plicata) again assuming the river form (costata). This case should be compared with what we have learned about Fusconaia ftava (See p. 25).
Genus Quadrula Rafinesque (1820).
Ortmann, 1912, p. 250; Simpson, 1914, p. 811.
Type Obliquaria metdnevra Rafinesque.
Five well-defined species and one variety are found in Pennsylvania.
Key to the Species and Varieties of Quadrula.
« !■ Shell more or less regularly rounded, with tubercles, which are rather small and irregularly scattered, ' and have no connection with the weakly developed, concentric beak-sculpture. Epidermis
yellow to brown, rajmd, when young, rays green, often very broad and broken up into large spots.
hi. Shell rounded, high, not transverse. Posterior wing not developed Q. pushilosa.
hi. Shell more transverse. Posterior wing better developed Q. pustulosa schooler aftensis.
ai. Shell not rounded, but trapezoidal, triangular, or elongated. Sculpture consisting of tubercles, knobs, or ribs, of a more or less definite arrangement, more or less related to the beak-sculpture; the latter double looped or of the zig-zag type. Epidermis rayed in various patterns or spotted. hi. Shell subtrapezoidal, with a narrow and blunt posterior ridge. In front of the latter a broad and shallow groove. Tubercles of posterior ridge not very large. Epidermis with ir- regular rays, rays often spread out, but not spotted.
Cl. Shell about as long as high. Groove of disk distinct, generally without nodules.
Q. quadrula.
34
MEMOIRS OP THE CARNEGIE MUSEUM.
C2. Shell much longer than high. Groove of disk shallow, and covered with the same nodules
as the anterior part of the shell Q. verrucosa.
62. Shell subtrapezoidal, with a broad and distinct posterior ridge. In front of this ridge, and behind, a slight depression, but no distinct radial groove. Tubercles of this ridge generally large. Rays of epidermis generally broken up into characteristic spots.
Cl. Shell subtrapezoidal or subrhomboidal, about as high as long. Sculpture of disk generally
well-developed Q. metanevra.
C2. Shell subrectangular, much longer than high. Sculpture of disk often poorly developed.
Q. cylindrica.
Quadrula pustulosa (Lea) (1831).
Quadrula pustulosa (Lea) Simpson, 1914, p. 848.
Plate III, figs. 4, 5.
Records from Pennsylvania :
Ilarn, 1891 (Western Pennsylvania).
Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver River, Wampum, Lawrence Co.). Ortmann, 19096, p. 199.
Characters of shell: Shell of not more than medium size, but rather heavy. Outline subcircular, or obovate, about as high as long, not oblique, but rather upright. Beaks moderately prominent. Beak-sculpture consisting of two to three indistinct, concentric ridges, slightly angled and nodulous behind, not con- tinued upon the disk. Shell from rather swollen to rather fiat, evenly convex, without, or with very indistinct, posterior ridge. Basal margin evenly rounded. Surface sculptured by very irregular and variable nodules or pustules, absent towards the beaks. Sometimes the surface remains entirely smooth, but generally the pustules begin at a certain distance from the beaks, and are larger or smaller, rounded ©r transverse. Upon the posterior slope they are absent or present, and, when present, generally smaller, and often arranged in radiating ribs, or they are entirely rib-like. Epidermis yellowish brown to brown, when young, generally with distinct green rays, one of which is characteristically broad, sometimes inter- rupted so as to form blotches. Growth-rests dark browm.
Hinge-teeth heavy, more or less ragged. Pseudocardinals slightly divergent. Interdentum present, narrower or broader. Lateral teeth rather strong and short. Beak cavity rather deep. Dorsal muscle scars upon the hinge plate. Nacre always white. No difference whatever between the male and female shell.
L. H. D.
Size: Kelly, Cat. No. 61.4365 (largest at hand, d') 72 mm. 70 mm. 36 mm.
Call (1896a, p. 43) says that “ the female is often somewhat emarginate.” This is not always so in the females at hand; in fact this slight emargination of the posterior margin, which is actually often observed in this species, has no relation to sex.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
35
Soft parts (See Ortmann, 1912, p. 251). Glochidia figured by Lefevre & Curtis (1910, p. 97, fig. F, and 1912, p. 146, fig. F), 0.23 X 0.32 mm.; by Surber (1912, PI. 2, fig. 20) 0.23 X 0.29 mm., and by Howard (1914, PL 5, fig. 36). I have seen them in specimens from West Virginia and Arkansas, and found them to vary somewhat in size: 0.21 X 0.26 mm. (Arkansas), and 0.22 X 0.29 mm. (West Virginia). They are rather large for the genus.
Breeding season: Gravid females have never been observed in our state, but I have them from Arkansas, collected on May 19, 1911, and July 19, 1913, and from West Virginia, collected July 10, 1911. In the upper Tennessee region, I found them on Alay 25, 1914. This is probably a summer breeder (tachytictic).
Remarks: This species is well-distinguished by the general shape and sculp- ture, although the latter is often poorly developed and even absent (chiefly in young specimens). The sculpture always begins at a certain distance from the beaks, and does not represent a continuation of the beak-sculpture. In its general character, this sculpture resembles that of certain species of other genera, chiefly of Plethobasus cooperianus (Lea), and there are cases, where it is hard to distinguish these two species by the shell alone. However, P. cooperianus is generally more oblique, and the nodules are rather distinctly confined to the posterior half of the shell. Of course, they are easily told apart, when the soft parts are at hand.
The shells from the Ohio-drainage in Pennsylvania are undoubtedly all typical Q. pustulosa. But outside of our state this species varies greatly, not only in sculpture, but also in shape. These variations are in part geographical, and have been named, but cannot be discussed here.
Localities in Pennsylvania, represented in the Carnegie Museum:
Ohio River, Shippingport, Cooks Ferry and Industry, Beaver Co.; Neville Island, Allegheny Co.
Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).
Mahoning River, Mahoningtown and Edinburg, Lawrence Co.
Monongahela River, Westmoreland Co., and Charleroi, Washington Co. (G. A. Ehrmann).
Cheat River, Cheat Haven, Fayette Co.
Allegheny River, Natrona, Allegheny Co.; Godfrey and Kelly, Armstrong Co.
Locality in Pennsylvania, represented in the Philadelphia Academy of Natural Sciences:
Ohio River, Coraopolis, Allegheny Co. (S..N. Rhoads).
Other localities, represented in the Carnegie Museum:
Ohio-Mississippi-drainage :
Ohio River, Toronto, Jefferson Co., Ohio; St. Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio.
Little Kanawha River, Grantsville, Calhoun Co. (F. W, Graham); BurngviUe, Braxton Co., West Vir- ginia.
36
MEMOIRS OF THE CARNEGIE MUSEUM.
Elk River, Shelton, Clay Co., West Virginia.
Pocatalico River, Rajnnond City, Putnam Co., West Virginia.
Little Coal River, Boone Co., West 'Virginia (Hartman collection).^®
Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.
Licking River, Farmer, Rowan Co., Kentucky.
Wabash River, New Harmony, Posey Co., Indiana (A. A. Hinkley).
Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection); and Moline, Rock Island Co., Illinois (P. E. Noordgrcn).™
Tennessee-drainage:
Tennessee River, Florence, Lauderdale Co., and Tuscumbia, Colbert Co. (H. H. Smith); Bear Creek, Burleson, Franklin Co. (H. H. Smith); Shoals Creek, Lauderdale Co. (H. H. Smith); Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).
Tennessee River, Concord and Knoxville, Knox Co., Tennessee.
French Broad River, Boyd Creek, Sevier Co., Tennessee.
Nolichucky River, Chunns Shoals, Hamblen Co., Tennessee.
Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co., Tennessee.
Clinch River, Solway, Knox Co.; Edgemoor, Clinton, and Offutt, Anderson Co., Tennessee.
Specimens from the Black River and Ouachita River in Arkansas do not represent the typical phase of Q. pustulosa: they require further study. (See; Wheeler, 1918, p. 12.3.)
a Quadrula pustulosa.
A Quadnda pustulosa schoolcraftensis.
• Quadrula quadrula.
+ Quadnda verrucosa.
Labeled “ Little Coal River, Logan Co., Virginia,” but this river does not flow through Logan Co. The specimens from the Mississippi River differ somewhat from the typical form in having a better developed posterior wing and somewhat more prominent beaks, so as to intergrade toward Q. dorfeuillana (Lea),
OETMANN: monograph of the naiades of PENNSYLVANIA.
37
Distribution and Ecology in Pennsylvania (See fig. 5) : This is one of the rarer species in Pennsylvania. It occurs in the Ohio and Monongahela, in the latter as far up as the lower Cheat River, but it is missing in the headwaters of this system in West Virginia. In the Allegheny River, it has been traced up into Armstrong Co., and, in addition, it is found in the Beaver and Mahoning Rivers, but has not been found in the Shenango.
In all the other streams, not mentioned, it is absent, and thus its ecological preferences are decidedly with the larger streams; but since it has not often been found alive, details are scarce. I collected it myself in the heavy gravel of the Ohio, Allegheny, and Cheat, as well as in coarse gravel in the Mahoning. On similar bottom I found it in the Little Kanawha and Elk Rivers in West Virginia, and in the upper Tennessee region. But in Pocatalico River I found it in pure sand. In the Ohio in West Vir*ginia and Ohio, it is abundant on the “ shell banks,” in deep, strongly flowing water, with gravelly bottom, and here it is frecpiently taken by the clam-diggers. Call (1900, p. 488) reports this species from muddy bottoms as well as from sand and gravel bars.
General distribution: Type-locality, Ohio (Lea). This species is quite char- acteristic of the Ohio proper and its larger tributaries. It also belongs to the Cumberland and Tennessee-drainages, and is there in its typical phase, but some confusion exists as to this.^^ In the states of Ohio, Indiana, and Illinois it is a common shell (Sterki, 1907a, Call, 1896a, and 1900, Baker, 1898a and 1906). How far it extends West and Southwest is hard to say, but the typical form has been figured by Scammon (1906) from Kansas. In this region, however, it is in part represented by more or less distinct varieties. Among the material at hand, there seem to be several of the latter, but I am well acquainted only with one (var. schoolcraftensis, see below) .
Note: This species is only moderately swollen, or even rather flat, in Penn- sylvania. Farther down the Ohio more distinctly swollen individuals are met with, becoming sometimes almost globular, culminating finally in the type known as dorfeuillana Lea (with high beaks). This indicates a tendency similar to that observed in several of the preceding species, which, however, is in this case not so distinctly marked, since Q. pustulosa does not go up, as a rule, into small streams.
The chief mistake has been made by Simpson (1900, p. 780) in stating that the varieties pernodosa Lea and kieneriana Lea (not keineriana ) are found in streams “ draining into the Gulf of Mexico,” while the type-locality of pernodosa, at least, is in the upper Tennessee drainage. A singular double mistake is made by Sterki (1907a, p. 391) in quoting “ kleineriana ” (for kieneriana) from Lake Erie.
38
MEMOIRS OF THE CARNEGIE MUSEUM.
Quadrula pustulosa schoolcraftensis (Lea) (1834),
Quadrula pustulosa schoolcraftensis (Lea) Simpson, 1914, p. 850.^^
Plate III, figs. 6, 7.
Never before reported from Pennsylvania.
Characters of variety: Distinguished from typical pustulosa by the more trans- verse and subquadrate outline. This is brought about by a stronger development of the posterior wing, and the shell thus appears more elongated, and less high. In addition, the posterior ridge appears more distinct, the color of the epidermis is generally lighter, and the nodules are rather poorly developed, but the latter characters are inconstant.
Size: 1. Erie, Pennsylvania, Cat. No. 61.4515 (old $ , much L. H. D.
elongated) 72 mm. 52 mm. 36 mm.
2. Cedar Point, Ohio, Cat. No. 61.4451 (normal). . ..49 “ 41 “ 26 “
Soft parts agreeing with those of the typical form (Ortmann, 1912, p. 251) Glochidia and breeding season not observed.
Remarks: The characteristic feature of this form, the transverse shape, has been sufficiently emphasized by the earlier authors, but later on it was uncon- ditionally thrown together with pustidosa (Call, 1900; Simpson, 1900). Sterki (1907a, p. 291), however, has again correctly recognized it, and Simpson follows him (1914). I also believe that it is a distinguishable form, but there are inter- grades with the normal pustulosa. In our region, schoolcraftensis is quite distinct, and does not come together geographically with the typical form. But this is the case elsewhere. Baker (1898a, PI. 24 and 25) has given a number of figures, some of which are evidently pustulosa, but at least one is an undoubted school- craftensis. There are some figures on Baker’s plates, which are more or less tran- sitional between the two, and such specimens are also represented in the Carnegie Museum. An example is a fine large specimen from Kishwaukee River, Rockford, Winnebago Co., Illinois (P. E. Nordgren) (L. 64, H. 60, D. 43 mm.). In outline this specimen is intermediate, but on the other hand it is rather smooth and has higher beaks, but not as high as in dorfeuillana. In the latter character it resembles some specimens from the Mississippi, taken at Moline, Rock Island Co., Illinois (P. E. Nordgren) and from Muscatine, Iowa, but the latter are not so transverse. (I have recorded these under pustulosa.)
Intergrades certainly exist, and as far as I can see at present, they are found in Illinois. They may exist elsewhere. The opinion of Call, Baker, Scammon,
Unio prasinus Conrad, made by Simpson a synonym of tj^pical Q. pustulosa, belongs here.
I
ORTMANN: monograph of the naiades of PENNSYLVANIA. 39
and others, that these forms belong to the same species, surely is founded upon the observation of transitions, and I also think that schooler aftensis should not be separated specifically, although the conditions in Pennsylvania would suggest such a step.
It should be mentioned that specimens from Lake Erie show the regular and distinct growth-lines, commonly observed in shells from Lake Erie.
Localities represented in the Carnegie Museum:
Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania.
Lake Erie, Cedar Point, Erie Co., Ohio (C. Brookover).
Lake Erie, La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).
Miami and Erie Canal, Waterville, Lucas Co., Ohio (C. Goodrich).
Maumee River, Defiance, Defiance Co., Ohio (C. Goodrich).
Grand River, Grand Rapids, Kent Co., Michigan (G. H. Clapp donor).
Coon River, Dallas Co., Iowa (Smith collection).
Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).
Hinkston Creek, Columbia, Boone Co., Missouri (D. K. Greger).
Platte River, Garretsburg, Buchanan Co., Missouri (W. I. Utterback).
Kansas and Wakarusa Rivers, Lawrence, Douglas Co., Kansas (R. L. Moodie).“
Chikaskia River, Tonkawa, Kay Co., Oklahoma (E. B. Isely).
In addition, there is a fine large specimen, labeled Poland, Mahoning Co., Ohio, and two others, labeled: Grand River, Ohio, all from the Hartman collection. Those from the latter locality were named dorfeuillanus, with the remark: “ Lea datum,” and thus they are supposed to be authentic specimens from Lea. But I believe, that these localities are unreliable. These specimens are so typically repre- sentative of the form from Lake Erie (with very distinct and regular growth-lines) that I think they are undoubtedly from the lake. The fauna of Grand River in Ohio is practically unknown. Poland is near the Mahoning River, and we know, that the typical pustulosa is found in the Pennsylvanian part of this river.^^
Distribution and Ecology in Pennsylvania (See fig. 5) : In Pennsylvania, this form has been found only in Presque Isle Bay of Lake Erie, in the characteristic fine sand of the “ flats ” and the North shore (“ Big Bend ”) of the bay, where the shore is lined with the Juncus americanus formation, in one to two feet of water. It appears to be a rare shell there, since only two specimens have turned up.
General Distribution: Type locality, “Fox River of Green Bay” in Wisconsin (Lea) . (This belongs to the Lake Michigan-drainage, and is at the same time the most northern locality known.)
From Lake Erie, this form has been reported before as Q. pustulosa (Walker, 1913). Aside from the localities given above for the lake, it is found in some of
Some of these specimens, when received, were labeled (by Scammon?) Quadrula rubiginosal Dean (1890) does not report pusiulosus from the Mahoning in Ohio, nor does Sterki (1907a), but both have pustulatus Lea, which is a species not at all found in this region. Probably, Dean’s pustu- latus is only a slip of the pen for pustulosus.
40
MEMOIRS OF THE CARNEGIE MUSEUM.
the tributaries in Ohio (Tiffin River, Sterki, 1907a, and other places in the Maumee- drainage), and is known from the Kankakee in Indiana (Sterki),®^ from Michigan, Illinois, and Iowa (Sterki, 1907a, Walker, 1894, as schoolcraftensis, and 1898 as pustulosa). That it is found in Iowa, is confirmed by the locality given above, and by Call (1895), while Geiser (1910) reports only pustulosa from the Wap- sipinicon River in northeastern Iowa. The siiecimens from Missouri and Kansas are fine and typical. Beyond this we do not know much about the distribution of this form, but it is apparent that this variety is more western and northern in its range than the typical pustulosa.
It should be noted that the form from Lake Erie hardly differs from the normal type of schoolcraftensis, except for the more regular and distinct growth-lines. The route by which it migrated into the lake is indicated by its presence in the Maumee-drainage.
(^UADRULA QUADRULA ( Rafiiiesciue) (1820).
Quadrula lachrymosa (Lea) Simpson, 1914, j). 841; Quaclrula quadrula (Rafin- esque) Vanatta, 1915, p. 556; Quadrula quadrula (Rafinesque) Utter- back, 1916, p. 53.
Plate IV, fig. I.
Records from Pennsylvania:
Ortmann, 19096, p. 199.
Characters of the Shell: Shell growing to a rather considerable size, rather heavy. Outline subtraiiezoidal or subquadrate, not oblique, not longer, or only slightly longer than high. Beaks moderately prominent, beak-sculpture double- looped, posterior loop slightly tubercular upon the iiosterior ridge, these tubercles continued upon the ridge as larger or smaller nodules. Anterior loop, toward the disk, breaking up into nodules, and this sculpture is continued upon the disk in an irregular way. Shell moderately swollen or rather flat, with a rather distinct, but narrow, posterior ridge. In front of the latter, there is a broad, shallow furrow, which is generally smooth, without tubercles. There are large nodules upon the posterior ridge, and smaller ones upon the disk in front of the furrow, but the most anterior part of the shell is generally smooth. Nodules and tubercles very varying in number, arrangement, size, and shape; often they are tear-like (verti- cally elongated), but they may be transversely elongated. The nodules of the anterior part of the shell are the direct continuation of the beak-sculpture. Pos- terior slope with more or less distinct nodules, often arranged in radiating ridges.
Specimens from the Kankakee River are cited as pustulosa by Wilson & Clark (1912a), but they mention the great variability.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
41
Epidermis yellowish to brownish, when young with indistinct rays, and the green color of the rays often spreads out over the shell in indistinct transverse bands or patches (under the larger nodules), so that the whole surface often appears more or less suffused with green. But there are never triangular spots (as in Q. metanevra and Q. cylindrica). Growth-rests more or less distinct, brownish. Old shells often are uniformly brown, and the sculpture disappears toward the lower margin.
Hinge-teeth moderately strong. Pseudocardinals divergent, ragged; inter- dentum moderately developed. Beak-cavity moderately deep. Dorsal muscle- scars on hinge-plate. Nacre white. No differences between the male and female shell.
L. H. D.
Size: I. Cooks Ferry, Cat. No. 61.3869 72 mm. 60 mm. 36 mm.
2. Erie, Cat. No. 61.4526 64 “ 52 “ 29 “
This species grows considerably larger outside of Pennsylvania.
Soft parts: Ortmann (1912, p. 253). Glochidia, figured by Surber (1915, p. 8, pi. 1, fig. 11), remarkably small (0.085 X 0.090 mm.), much resembling those of Q. verrucosa. Howard (1914, })1. 5, fig. 29) also gives a figure.
Breedmg season: Wilson & Clark (1912a, p. 43) report this species as found beginning to become gravid on August 21 in Illinois, and Surber found specimens with glochidia in August in Kansas.
• Remarks: The broad, smooth furrow of the disk and the sculpture distinguish this species sufficiently from all other Pennsylvanian forms. The sculpture is very variable, sometimes rather slightly developed, in other cases strong and very prominent. Only a few specimens having been found in our state, no important variations are to be noted. The color-pattern of this species, although quite variable, is characteristic in so far that the rays never break up into well-defined spots.
Localities in Pennsylvania, represented in the Carnegie Museum:
Ohio River, Cooks Ferry, Beaver Co.
Lake Erie, Presque Isle Bay, Erie, Erie Co.
Other localities represented in the Carnegie Museum:
Miami-Erie Canal, Lucas Co., Ohio (C. Goodrich).
Ohio River, St. Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co., West Virginia; Ports- mouth, Scioto Co., Ohio.
Middle Island Creek, Union Mills, Pleasants Co., West Virginia.
Wolfe Creek, Wolfe P. 0., Washington Co., Ohio (W. F. Graham).
West Fork White River, Riverside, Greene Co., Indiana (.1. D. Haseman).
Wabash River, New Harmony, Posey Co., Indiana (A. A. Hinkley).
42
MEMOIRS OF THE CARNEGIE MUSEUM.
Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).
Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection).
Sheyenne River, Argusville, Cass Co., North Dakota (S. M. Edwards).
Hinkston Creek, Columbia, Boone Co., Missouri (D. K. Greger).
Platte River, Garretsburg, Buchanan Co., Missouri (W. I. Utterback).
Lake Contrary, St. Joseph, Buchanan Co., Missouri (W. I. Utterback).®*'
Wakarusa River, Lawrence, Douglas Co., Kansas (R. L. Moodie).
Chikaskia River, Tonkawa, Kay Co.; North Fork Canadian River, Weleetka, Olcfuskee Co.; Deep
Fork Canadian River, Okmulgee, Okmulgee Co., Oklahoma (F. B. Isely).
Distribution and Ecology {See fig. 5). Type locality : Ohio River (Rafinesque) according to Vanatta, Salt River, Kentucky.
Two specimens only have been found in Pennsylvania; one in the Ohio River, just above the Ohio state line, the other in Lake Erie. The first was found in sandy-muddy bottom in a quiet eddy in a riffle, the second (a dead shell) in fine sand, the environment characteristic of Presque Isle Bay. According to Baker (1898a, p. 85) this species prefers the muddy and sandy bottoms of lakes and larger rivers, and Call (1900, p. 490) calls it a mud-inhabiting shell, while Scammon (1906, p. 252) says that in Kansas it is not particular as to station, but prefers sand. Apparently, it does not find congenial habitats in Pennsylvania, except in Lake Erie and in the Ohio. Its utmost upstream migration has barely reached our state.
Farther down the Ohio, it is also not very abundant, but it is present. In the slack water of the lower part of Middle Island Creek, I have seen a number of dead shells, and here it goes up as far as the slack water, about 5 miles, to just below Union Mills. Also farther down it is not abundant, and distinctly prefers sandy or muddy bottom to gravel. It does not ascend far into the tributaries, but it goes into the lower Muskingum and to Wolfe Creek. It is not in the Tus- carawas, according to Sterki (1907a, p. 390). In western Ohio and in southern and northwestern Indiana it is more widely distributed. Here it crosses over into the Lake Erie-drainage (Ohio Canal, Sterki), and into the lake (Sterki, and Walker, 1913). It occurs also in Beaver Creek, Lorain, Lorain Co., Ohio (Dali & Simpson, 1895), and in Michigan (Walker). It extends westwards and southwards to Minnesota, Iowa, Kansas, Oklahoma, northeastern Texas, and northern Louisiana. It also crosses over into the drainage of the Red River of the North (Winnipeg, Canada). There are no records from south of the Ohio, but the Carnegie Museum has it from the Tennessee in northern Alabama. From Alabama, there is only a single, doubtful and indefinite, previous record (Call).
Thus it seems that the center of this species is in the Mississippi and lower Ohio.
This is the var. contraryensis of Utterback, a local phase.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
43
Its absence in the Cumberland and upper Tennessee is remarkable, but probably accounted for by its ecological habits. On the Gulf plain, it is largely represented by other allied species.
Sterki says that the Lake Erie form is “little inflated and has few tubercles.” This fits the specimen before me, but I cannot judge from a single example whether this is a constant difference.
Quadrula verrucosa (Rafinesque) (1820).
Tritocgonia tuberculata (Barnes) Simpson, 1914, p. 318; Tritogonia verrucosa
(Rafinesque) Vanatta, 1915, p. 554; Quadrula verrucosa (Rafinesque)
Utterback, 1916, p. 62.
Plate IV, figs. 2, 3.
Records from Pennsylvania:
Rlioads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver River, Wampum, Lawrence Co.). Ortmann, 19096, p. 198.
Characters of Shell: Shell large and heavy. Outline elongate-sub trapezoidal, considerably longer than high. Beaks low. Beak-sculpture consisting of one or two subconcentric bars, curving upwardly and nodular behind, followed by addi- tional bars, which are more or less distinctly double-looped, and break up into nodules, which are indistinctly arranged in zig-zag waves. This sculpture is con- tinued upon the disk. Shell moderately swollen or rather flat, with a distinct, but narrow posterior ridge. In front of the latter the sides of the shell are flattened or somewhat concave, thus forming a broad, shallow radial groove. The posterior nodular part of the beak-sculpture is continued upon the posterior ridge as a row of tubercles, which, however, are not very prominent, and gradually disappear. In front of the posterior ridge the disk, including the groove, is thickly studded with rather small, low tubercles, which are often more or less distinctly arranged in diagonal rows, and represent the continuation of the beak-sculpture. Toward the lower margin, these tubercles become irregular, and may disappear. Posterior slope with ribs radiating from the posterior ridge; they are small, irregular, and nodulous toward the beaks, but become larger toward the posterior end of the shell. Epidermis yellowish to rusty brown and blackish, without distinct rays, but gen- erally suffused with green in irregular patches. Often the whole surface is greenish, and only the tubercles are more or less yellowish or brownish. Growth-rests more or less distinct. Old shells are generally uniformly dark brown or blackish.
Hinge-teeth well-developed, strong. Pseudocardinals divergent, large, ragged. Interdentum moderately developed. Lateral teeth long, straight, rather heavy.
44
MEMOIRS OF THE CARNEGIE MUSEUM.
Beak-cavity not very deep. Dorsal muscle-scars on the hinge-plate. Nacre white.®^
In this species, we meet with a more or less distinct sexual difference in the shell, unusual in the subfamily Unionince. In the male the posterior margin of the shell is short and subtruncate, and the ribs of the posterior slope are generally well-developed and continued to the margin. In the female the posterior margin is broadened and flattened in the region of the posterior angle, and the ribs of the posterior slope are poorly developed and broadened, so that the posterior end of the shell appears expanded. This expansion corresponds to the location of the
anal opening in the soft parts.
L. H. D.
Size: 1. Edinburg, Cat. No. 61.3577 (sex doubtful) 156 mm. 86 mm. 43 mm.
2. Mahoningtown, Cat. No. 61.3574 (cf) 121 “ 70 “ 44 “
3. Pulaski, Cat. No. 61.4548 (9) 130 “ 64 “ 37 “
No. 1 is the largest specimen from Pennsylvania, but larger ones are known from elsewhere. Scammon (1906) gives 168 as maximum length.
Soft parts (See Ortmann, 1912, p. 254). The glochidia have been figured by Surber (1912, PI. 2, fig. 31). They are unusually small, 0.085 X 0.100 mm.
Breeding season: Sterki (19076, p. 48) collected gravid females on June 10,
1907, and Surber gives April to June. I did not find any gravid females in Penn-
«
sylvania, but collected some on May 9, 1913, in Pocatalico River, and on May 23, 1912, in West Fork River, West Virginia. All these had eggs only, filling all four gills, forming lanceolate, not very solid, white placentae. Those from , Poca- talico River (two specimens) had the eggs brownish, discolored, and they were being discharged when opened (about two hours after capture). This was appa- rently premature discharge, and the eggs were dead (suffocated).
Remarks: This species has a very characteristic outline and sculpture. The latter, however, is quite variable. The tubercles of the disk may be larger or smaller, regular or irregular in size, rounded, subtriangular, or even vertically elongated. The ribs of the posterior slope are also variable, and more or less distinct. Toward the lower margin, the sculpture often disappears, and in some individuals the whole lower half of the shell may be smooth. The most remarkable fact in this species is the sexual differentiation of the shell, which has led Simpson to the opinion that it should represent a separate genus {Tritogonia) . However, as I have shown, in all other characters, chiefly those of the soft parts, it is a true Quadrula. The posterior dilatation of the shell in the female is in an entirely different region from that in the females of the Lampsilince.
In Pennsylvania, the nacre is always white; in the South (Alabama, etc.) it may be purple.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
45
The sexual differences are sometimes quite striking, but not always so. It is by no means ^‘one of the easiest species in which to distinguish the sexes,” as Scammon (1906, p. 314) says. In the largest specimen, mentioned above, I am in fact in doubt whether it is a male or a female, and it is true that sometimes in large males (determined by the soft parts), the posterior end of the shell is some- what expanded, with less developed ribs. But, as a rule, in individuals of medium and large size, the sex may be easily told from the shape of the shell. In young shells this is very hard, or even impossible to do.
It should be mentioned that the fem.ale shell is on the average more flattened and compressed than that of the male. This also is a rather unusual feature.
Localities in Pennsylvania, represented in the Carnegie Museum:
Ohio River, Industry, Beaver Co.
Beaver River, 'Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).
Mahoning River, Mahoningtown, Coverts, and Edinburg, Lawrence Co.
Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsville, Mercer Co.
Pymatuning Creek, Pymatuning Township, Mercer Co.
Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).
Dunkard Creek, Wiley and Mount Morris, Greene Co.
Allegheny River, Kelly, Armstrong Co.
Locality in Pennsylvania, represented in the Philadelphia Academy of Natural Sciences:
Ohio River, Coraopolis, Allegheny Co. (S. N. Rhoads).
Other localities represented in the Carnegie Museum:
Ohio-drainage :
Tuscarawas River, Ohio (Holland collection).
Wolfe Creek, Washington Co., Ohio (W. E. Graham).
West Fork White River, Riverside, Greene Co., Indiana (J. D. Haseman).
West Fork River, Lynch Mines, Harrison Co.; WTst Milford, Harrison Co. (W. F. Graham); Lightburn, Lewis Co., West Virginia.
Little Kanawha River, Burnsville, Braxton Co., West Virginia.
Pocatalico River, Raymond City, Putnam Co., West Virginia.
Elk River, Sutton and Gassaway, Braxton Co., West Virginia.
New River, Hinton, Summers Co., West Virginia.
Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.
Licking River, Farmer, Rowan Co., Kentucky.
Tennessee-drainage :
Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).
Shoals Creek, Lauderdale Co., Alabama (H. H. Smith); Elk River, Fayetteville, Lincoln Co., Tennessee (H. H. Smith); Hurricane Creek, Gurley, Madison Co., Alabama (H. E. Wheeler); Paint Rock River, Trenton and Princeton, Jackson Co., Alabama (H. H. Smith); Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).
46
MEMOIRS OF THE CARNEGIE MUSEUM.
^Yest of Mississippi:
Meramec River, Meramec Highlands,, St. Louis Co., Missouri (N. M. Grier).
Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).
White River, Cotter and Norfolk, Baxter Co., Arkansas (A. A. Hinkley).
Saline River, Benton, Saline Co., Arkansas (H. E. Wheeler).
Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).
Illinois River, Talequah, Cherokee Co., Oklahoma (F. B. Isely).
Bayou Pierre, De Soto Parish, Louisiana (L. S. Frierson).
Mississippi, Alabama, Georgia:
Pearl River, Mississippi (Juny collection).
Buttahatchee River, Hamilton, Marion Co., Alabama (H. H. Smith).
Sipsey River, Texas, Marion Co.; Fayette, Fayette Co.; and Elrod, Tuscaloosa Co., Alabama (H. H. Smith).
Forks of Black Warrior River, Walker Co., and Black Warrior River, Squaw Shoals, Jefferson Co., Alabama (H. H. Smith).
Valley Creek, Toadvine, Jefferson Co., Alabama (H. H. Smith).
North River, Haglers Mill, Tuscaloosa Co., Alabama (H. H. Smith).
Cub Creek, Pine Hill, Wilcox Co., Alabama (H. H. Smith).
Cahaba River, Gurnee, Shelby Co., Alabama (H. H. Smith).
Coosa River, Wetumpka, Elmore Co.; Weduska Shoals, Shelby Co.; Riverside, St. Clair Co., Alabama (H. H. Smith).
Choccolocco Creek, Calhoun Co., Alabama (H. H. Smith).
Oostanaula River, Rome, Floyd Co., Georgia (G. H. Clapp donor).
Conasauga River, Whitfield Co., Georgia (H. H. Smith).
Distribution and Ecology in Pennsylvania (See fig. 5) : In Pennsylvania this species is rather local and not widely distributed. In the Ohio, Monongahela, and Allegheny, it is rather scarce, and only isolated specimens turn up. In the drainage of the Beaver it is a little more frequent, and the best places are at Edin- burg and Pulaski. This would indicate, that the species prefers smaller rivers, which is substantiated by the fact that it is rather abundant in the headwaters of the Monongahela, in Dunkard Creek in Pennsylvania, and in West Fork River in West Virginia. In the upper Little Kanawha and in Elk River, West Virginia, it is also frequent.
This shell is found in coarse or fine gravel, generally not very deeply buried, but young shells are often found in fine sand and even in mud. In a mill-race of the Mahoning, at Edinburg, among heavy rocks, about a dozen were collected, all very large and ponderous. The best places are riffles with strong currents, interrupted by patches of Dianthera americana.
Scammon (1906, p. 314) reports this species in various ecological surroundings, but says it prefers gravel and shingle, with a swift current, while Call (1900, p. 465) says that it delights in muddy bottoms, which is surely not the case in Penn- sylvania.
OETMANN; monograph of the naiades of PENNSYLVANIA.
47
General distribution : Type locality, Ohio River (Rafinesque) .
Simpson (1900) gives as the range of this species: Mississippi-drainage generally; streams falling into the Gulf of Mexico from the Alabama system west to central Texas.” This indicates a very wide distribution. In the Mississippi it goes into Wisconsin (Barnes, 1823; Lapham, 1860), and southern Minnesota (Grant, 1886; Holzinger, 1888), but its northward extension is generally rather restricted. Call (1895, p. 55) reports it from western New York, which is un- confirmed, and is very questionable, judging from its absence in the headwaters of the Allegheny in Pennsylvania, and its entire absence from the whole lake-drainage in Pennsylvania, Ohio (Sterki, 1907a), Indiana (Call, 1896a) and Illinois (Baker, 1906). As our records show, it occurs in the tributaries of the Ohio in West Vir- ginia and Kentucky, and goes up the Tennessee to northern Alabama, but is absent from the upper Tennessee region, except the Hiwassee River. Westward it goes to Iowa, Kansas, and Oklahoma, and its occurrence in Arkansas, and in the Gulf-drainage from Georgia to Texas is well established.
This species seems to belong more to the south and the west, but it has as- cended the Mississippi to a considerable distance, and the Ohio practically through- out its whole drainage, except the smallest headwaters in the north and east.
It does not seem to be very abundant in the larger streams. I never found it in the Ohio proper below the Pennsylvania state-line, although it is in the Cin- cinnati list of shells.
Quadrula metanevra (Rafinesque) (1820).
Quadrula metanevra (Rafinesque) Simpson, 1914, p. 834.
Plate IV, figs. 4, 5, 6.
Records from Pennsylvania:
Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.).
Ortmann, 19096, p. 198.
Characters of the shell: Shell of medium size, heavy. Outline subtrapezoidal, subrhomboidal, or subquadrate, sometimes subtriangular, short, not longer, or very little longer, than high. Beaks moderately elevated. Beak-sculpture con- sisting of two or three subconcentric bars, which are angular and nodulous upon the posterior ridge; anterior part rather straight, and curved upward in front, thick, but not sharply marked. The nodulous portion is repeated upon the posterior ridge in the shape of tubercles, while the anterior part of the following bars becomes indistinct, and is soon supplanted by the sculpture of the disk. Occasionally the third and fourth bars show an indication of double-looped structure, and sometimes
48
MEMOIRS OF THE CARNEGIE MUSEUM.
even traces of a breaking up into small zig-zag nodes is seen. In certain other cases the beak-sculpture is separated from that of the disk by a smooth space. Shell sometimes rather flat, but generally more or less swollen, with a broad, dis- tinct, and prominent posterior ridge. In front of the ridge, the shell is flat or dejiressed, but without forming a distinct radiating furrow. The posterior nodular l)art of the beak-sculpture is continued upon the ridge as large and distinct promi- nent tubercles or nodes, standing rather remote from each other; sometimes these nodes are broken uj) into clusters of nodules, and in rare cases they are rudi- mentary or absent. In front of the posterior ridge the shell is covered by larger or smaller nodules, very irregular and variable in arrangement, shape, and number. Sometimes there are very few, in other cases very many of them, but toward the lower margin they generally tend to disappear in larger shells, and the anterior jiart of the disk is always smooth. Towards the beaks, the nodules are smaller, and may disaiipear, or pass into the beak-sculpture. Posterior slope generally marked off from the posterior ridge by a narrow furrow, iiroducing an emargination of the iiosterior margin, and its surface may be smooth, or tuberculated, or radiately ribbed. Epidermis yellowish to brownish black, with a characteristic greenish color-])attern : the raja's are broken up into triangular spots of larger or smaller size, pointed toward the lower margin. These color-patches are very variable, sometimes almost absent, sometimes very distinct, chiefly so in young specimens; in older ones they become obliterated, the epidermis appearing uniformly brownish. Regular linear ra3^s are sometimes present, but not often. Growth-lines more or less distinct.
Hinge-teeth well-developed, strong. Pseudocardinals divergent, large, ragged. Interdentum jiresent, generall}^ well-developed. Lateral teeth strong, of medium length. Beak-cavity deep or moderately' deep. Dorsal muscle-scars on the hinge-plate. Nacre always white.
Sexes absolutely indistinguishable in the shell.
Size: 1. Industry, Cat. No. 61.3866 (largest from L. H. D.
Pennsylvania) 90 mm. 71 mm. 47 mm.
2. Kelly, Cat. No. 61.3861 (very flat) .89 “ 70 “ 36 “
3. Aladdin, Cat. No. 61.3868 (normal) 67 “ 58 “ 38 “
Soft parts (See Ortmann, 1912, p. 255, fig. 6). Glochidia figured by Lefevre & Curtis (1910, }). 97, fig. E, and 1912, p. 146, fig. E), by Surber (1912, PI. 2, fig. 26), and Howard (1914, PL 5, fig. 31). Their measurements have been given as 0.18 X 0.19 mm. and 0.175 X 0.200 mm. I have not seen glochidia in Pennsyl- vania, but have found them in the flat form (var. wardi) from the Little Kanawha
ORTMANN: monograph of the naiades of PENNSYLVANIA.
49
River, West Virginia. The shape was the same, as reported, lint they were slightly larger: 0.20 X 0.22 mm. I have repeatedly found specimens with eggs and with distinct placentae; but when giochidia are present, the placentae easily fall apart.
Breeding season: The specimen with giochidia was found on May 24, 1911, and a specimen with eggs was collected on June 22, 1909. Other specimens with eggs were received from Arkansas, collected on June 26, 1911. The early date for giochidia is remarkable. It is astonishing, on the other hand, that among numerous specimens collected on June 20, 21, and 22, 1911, in the Ohio River at St. Maiys, West Virginia, no gravid females were found. The species surely is tachytictic; Surber (1912, p. 7) gives May to July as the breeding season.
Remarks: It is hardly possible to mistake this species on account of its marked shape and sculpture. Yet there is quite a range of variation, chiefly with regard to obesity of the shell, and to sculpture. As is seen in the measurements given above, some shells are considerably more flattened, and it is to be noted that such specimens are more frequent in the Allegheny than in the Ohio. This would express the same tendency toward flattening of the shell in smaller rivers, which we have noticed in several of the foregoing species. But in this case it is impossible to distinguish a flat race, since there are all intergrades, and only few flat indi- viduals are found, always associated with others which are swollen (See PI. IV, figs. 4, 5, 6).
In the development of the tubercles we notice an important variation in which the large tubercles of the posterior ridge are obliterated. Such specimens, which at the same time were rather flat, have been described by Lea (Obs. IX, 1863) as a separate species, Unio wardi (from Walhonding, Ohio; Wapsipinicon, Iowa; and Coal River, West Virginia).’* Three specimens from the latter locality are in the Carnegie Museum, derived from the Hartman collection. They undoubtedly are the same specimens referred to by Lea, as being in the Hartman collection. They agree very well with Lea’s figure, but the posterior ridge is not entirely smooth, although without the large tubercles. I have specimens like these from the Little Kanawha River and from the West Fork River. These also are remark- ably flat. Finally specimens resembling these are sometimes found in western Pennsylvania (See PI. IV, fig. 6). The second one, of which the measurements are given (from Kelly), should be by all means regarded as wardi and there is a half shell from the Monongahela at Charleroi, which has been labeled by Simpson as var. wardi, and rightly so. Compressed specimens, which also have the tubercles more or less obliterated, have been found occasionally all the way down from the
Lea says: “ Coal River, Logan Co., Va.,” but Coal River and Little Coal River are today in Kanawha and Boone Cos., West Virginia.
50
MEMOIRS OF THE CARNEGIE MUSEUM.
Allegheny, Armstrong Co., Pennsylvania to Parkersburg, West Virginia, but in company with the normal form. They should be called wardi, but do not form a distinct race, but are rather individual variations.
It is interesting to note, that outside of our state, nearly all specimens from small rivers and creeks, such as West Fork River, the Little Kanawha, and Coal
■ Quadrula metanevra.
• Quadrula cylindrica.
River, belong to this form called wardi, and it may be, that in such small streams, the wardi-type becomes the prevailing one, thus forming an ecological race. Sterki (1907a, p. 390) mentions such a case from Sugar Creek, a small tributary of the Tuscarawas River in Ohio.
Localities in Pennsylvania, represented in the Carnegie Museum:
Ohio River, Shipping-port, Cooks Ferry, and Industry, Beaver Co.; Coraopolis (S. N. Rhoads) and Neville Island, Allegheny Co.. «
Allegheny River, Aladdin, Godfrey, Johnetta, Kelly, and Templeton, Armstrong Co.
Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).
Locality in Pennsylvania, represented in the Philadelphia Academy of Natural Sciences:
Ohio River, Beaver, Beaver Co. (S. N. Rhoads).
Other localities represented in the Carnegie Museum:
Ohio-drainage:
Ohio River, Toronto, Jefferson Co., Ohio; Wheeling, Ohio Co., West Virginia (W. F. Graham); Claring- ton, Monroe Co., Ohio; St. Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio.
West Fork River, West Milford, Harrison Co., West Virginia (W. F. Graham).
ORTMANN: monograph of the naiades of PENNSYLVANIA.
51
Little Kanawha River, Grantsville, Calhoun Co. (W. F. Craham), and Burnsville, Braxton Co., West Virginia.
Coal River and Little Coal River, Boone Co., West Virginia (Hartman collection).^®
Tennessee-drainage :
Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith); Knoxville, Knox Co., Tennessee. Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).
Holston River, Mascot, Knox Co., Tennessee.
Mississippi-drainage and westward:
Peoria Lake, Illinois River, Illinois (Hartman collection).
Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection).
Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Crier).
Black Rock River and Spring River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).
Spring River, Williford, Sharp Co., Arkansas (H. E. Wheeler).^®
White River, Cotter, Baxter Co., Arkansas (A. A. Hinkley).
Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).
Neosho River, Miami, Ottawa Co., Oklahoma (F. B. Isely).
Alabama-drainage:
Sipsey River, Texas, Marion Co., Alabama (H. H. Smith).
Coosa River, Wilsonville, Shelbj^ Co., and Riverside, St. Clair Co., Alabama (H. H. Smith).
Distribution and Ecology in Pennsylvania (See fig. 6) : In Pennsylvania this species is restricted to the three large rivers, the Ohio, Allegheny, and Monongahela. In the Allegheny it goes up a little beyond the middle of Armstrong County. From the Monongahela it is known only from one locality, but it must have at one time ascended into West Virginia, for it occurs in the West Fork River.
Wherever found it is not rare, but it is most abundant in the Ohio below Pittsburgh. Here it favors the same places as most other species: coarse gravel in swiftly running water, and it helps to compose the shell-banks, where such are present (at Shippingport and Industry). This agrees with Scammon (1906, p. 350), who says that gravel-bars are its favorite habitat. In the Ohio below Wheel- ing it is frequently taken by the clam-diggers out of deep water in strong and steady currents.
General distribution: Type locality, Kentucky River (Rafinesque) .
The general distribution, as given by Simpson (1900) is: ‘‘Mississippi-drainage area, except its southern portion, extending to the Tennessee and Arkansas River.” This is quite correct, and it is to be particularly noted, that toward the east and north, this species does not cross over into any other drainage-system. As we have seen, in western Pennsylvania it does not advance far up into small streams.
See above p. 49. Specimens from West Fork, Little Kanawha, aud Coal Rivers all (together six) represent the form wardi.
This specimen is a good representative of the var. wardi,
52
MEMOIRS OF THE CARNEGIE MUSEUM.
and the same seems to be true elsewhere, although there are a few exceptions, and it seems that in such cases the main form tends to be replaced by the variety wardi.
In Ohio the main species hardly occurs outside of the Ohio proper (Sterki, 1907a). In Indiana it is known only from the larger rivers, the Ohio, White, Wabash, and Kankakee, and also from Eel, Blue, and Whitewater Rivers (Call, 1896 and 1900). In Illinois the same rule seems to prevail, so that its range covers the greater part of the state, except the northern and northeastern extremity (Baker, 1906; Wilson & Clark, 1912a). In the Mississippi-drainage, this species goes up into the Wisconsin River in Wisconsin, and the Minnesota River in Minne- sota.
Westward,s it is found as far as southeastern Kansas (Scammon, 1906) and to the southwest it extends to Oklahoma and the Ouachita River in Arkansas (Call, 1895; Wheeler, 1918, and our own material). The records from Louisiana and Texas are doubtful.
South of the Ohio in West Virginia this species is found in the headwaters of the Monongahela (West Fork River), in the Little Kanawha, and in Coal River, but, according to specimens in the Carnegie Museum, in the form wardi. Records from Kentucky and Tennessee are scanty. The type-locality is the Kentucky River. It is known from the Cumberland River (Scammon, 1906, and Wilson & Clark, 1914) and our material shows its presence in the Tennessee up to the lower Holston (See also Lewis, 1871). It turns up again in the Alabama system (Lewis, 1877, Call, 1885, Carnegie Museum). Lea’s locality (Obs. 10, 1863, p. 430), Columbus, Lowndes Co., Mississippi, belongs with this group of stations.
This latter set of localities is interesting, in view of Simpson’s statement, that it is missing in the lower Vlississippi region. If this is correct, it could have reached the Alabama-drainage only by crossing over from the Tennessee-drainage. This case should be kept in mind, for it is of zoogeographical importance.
Quadrula cylindrica (Say) (1817).'*^
Quadrula cylindrica (Say) Simpson, 1914, p. 832.
Plate V, figs. 1, 2, 3.
Records fro7n Pennsylvania:
Harn, 1891, p. 136 (western Pennsylvania).
Rhoads, 1899, p. 136 (Ohio River, Coraopolis, Allegheny Co.; Beaver, Beaver Co.; and Beaver River,
Wampum, Lawrence Co.).
Ortmann, 19095, p. 198.
Characters of the shell: Shell rather large, heavy. Outline elongated sub-
^ Not 1816 (See footnote 20 under Amhlema plicata, p. 25).
ORTMANN: monograph of the naiades of PENNSYLVANIA.
53
trapezoidal, or almost rectangular, considerably longer than high. Beaks moder- ately prominent. Beak-sculpture consisting of a number of bars, the first of which seems to be simple, the following two or three distinctly double-looped, with the posterior loop angular and nodose upon the posterior ridge. Farther on, upon the disk, the nodes of the posterior loop are continued a^s strong tubercles, while the anterior loop breaks up into small granules, which generally assume a zig- zag arrangement. They are continued downward to a variable extent, but gen- erally they soon disappear. Shell greatly swollen, from almost subcylindrical to rather flat, with a broad and distinct posterior ridge. In front of the ridge the shell is flattened, but has no radial furrow; behind the ridge the shell is more or less depressed, thus generally producing a slight emargination of the posterior margin. Upon the posterior ridge stands a row of tubercles (continuing the beak- sculpture), which increase in size toward the posterior angle, and may be larger or smaller; in some cases they are more or less obliterated. In front of the pos- terior ridge the shell is generally smooth, but towards the beaks we find a number of small tubercles or nodules, described above as a continuation of the broken-up anterior loops of the beak-sculpture. The development of these nodules is quite variable. The posterior slope may be ornamented with ridges or radiating rows of nodules, or may be entirely smooth. Epidermis yellowish to brownish or greenish, with dark green rays broken up into a characteristic pattern of triangular spots, pointed toward the lower margin. The distribution and size of these spots is rather irregular and variable, but they are generally well-developed and distinct, and only in rare cases do they become obliterated. Linear rays are, as a rule, entirely absent, indications of them are seldom seen upon the posterior ridge or posterior slope.
Hinge-teeth well-developed, but not very heavy. Pseudocardinals divergent, ragged. Interdentum narrow or almost absent. Lateral teeth long. Beak- cavity moderately deep. Dorsal muscle-scars on the hinge-plate. Nacre always white.
Sexes absolutely indistinguishable in the shell.
L. H. D.
Size: 1. Utica, Cat. No. 61.3852 (without any nodes) .... 123 mm. 50 mm.
2. Meadville, Cat. No. 61.3850 (without any nodes) .119 “ 54 “
3. Godfrey, Cat. No. 61.4358 (with strong nodes) . . 83 “ 33 “
4. Aladdin, Cat. No. 61.3856 (with strong nodes) . . 84 " 37 “
45 mm. 36 “ 33 “ 30 “
Pennsylvanian specimens compare favorably with previous measurements; in fact, the size of the first one given represents the maximum on record.
Soft parts (See Ortmann, 1912, p. 256). It is to be added that gravid females
54
MEMOIRS OF THE CARNEGIE MUSEUM.
have been observed subsequently. All four gills are used as marsupia. The eggs form placentie of lanceolate shape, of a peculiar yellow-brown or pale orange color. Glochidia have been observed in specimens from Holston River in Tennessee. Their shape is subcircular; length and height about 0.19 mm. Thus they resemble those of Q. metanevra (0.18 X 0.19), but they are more nearly circular.
Breeding season: Gravid females were found on May 22, 1914; May 23, 1914; May 24, 1911^ May 25, 1914; July 8, 1913. Glochidia were observed on May 23. The species apparently is bradytictic, as usual in the genus, but the date for glo- chidia is rather early. Wilson & Clark (1914) found gravid specimens in the Cumberland in June and July.
Remarks: This is an eminently characteristic sjiecies. The chief variations observed concern the general shape of the shell and the sculpture. The typical shape is “subcylindrical,” with the height about the same, or nearly the same, as the diameter (see measurements of Nos. 1 and 3, above), but there are rather flat individuals. Sterki (1907a, p. 390) calls attention to a form from the Tus- carawas River, which lacks the tubercles of the posterior ridge. This form is the lirevailing one in western Penng;ydvania, and is most abundant in the Beaver drainage and in French Creek. I have many specimens in which no trace of the tubercles is seen, and generally also the smaller nodules of the anterior part of the shell near the beaks and the sculpture of the posterior slope are absent in them, so that the shell appears absolutely smooth. Yet there are all sorts of intergrades, connecting this smooth with the normal form (See PI. V, figs. 2, 3). Farther down in the Ohio this smooth form is not found. These conditions are to a degree parallel with those observed in Q. jnetanevra and its var. wardi.
Very often the degree of obesity of the shell is correlated with the develop- ment of the sculpture, so that smooth specimens are at the same time unusually compressed. But this is not always the case, and smooth individuals may be subcylindrical, as our No. 1 (See measurements) and sculptured individuals may be somewhat compressed (See No. 4).
This smooth, and sometimes compressed form of the headwaters is so far known only from the Tuscarawas River in Ohio, from Beaver River and French Creek, in Pennsylvania, and from the upper Monongahela (West Fork River) in West Virginia.
It should be mentioned in this connection that a similar, compressed form is known from the headwaters of the Clinch River, in Virginia. This is the var. stngillata (Wright), but it is not a smooth form. On the contrary it is covered by a multitude of small tubercles. But here also the large tubercles of the posterior ridge are absent.
OETMANN: monograph of the naiades of PENNSYLVANIA.
Localities in Pennsylvania, ^represented in the Carnegie Museum:
Ohio River, Shippingport, Cooks Ferry, and Industry, Beaver Co.
Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).
Malioning River, Mahoningtown, Coverts, and Edinburg, Lawrence Co.
Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Clarksville and Shenango, Mercer Co. Pymatuning Creek, Pymatuning Township, Mercer Co.
Allegheny River, Aladdin, Godfrey, Johnetta, and Kelly, Armstrong Co.
French Creek, Utica, Venango Co.; Cochranton and Meadville, Crawford Co,
Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).
Localities in Pennsylvania, represented in the Philadelphia Academy of Natural Sciences:
Ohio River, Beaver, Beaver Co., and Coraopolis, Allegheny Co. (S. N. Rhoads).
Other localities represented in the Carnegie Museum:
Ohio-drainage :
Ohio River, Toronto, Jefferson Co., Ohio; Parkersburg, Wood Co., West Virginia; Portsmouth, Scioto Co., Ohio.
Tuscarawas River, Ohio (Holland collection).
West Fork River, West Milford, Harrison Co., W. Va. (W. F. Graham).^^
Little Kanawha River, Grantsville, Calhoun Co. (W. F. Graham), and Burnsville, Braxton Co., West Virginia.
Tennessee-drainage :
Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).
Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).
Paint Rock River, Paint Rock, Trenton, and Princeton, Jackson Co., Alabama (H. H. Smith).
Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill and Noeton, Grainger Co.; Austin Mill and Church Hill, Hawkins Co., Tennessee.
North Fork Holston River, Rotherwood, Hawkins Co., Tennessee.
Big Mocassin Creek, Mocassin Gap, Scott Co., Virginia.
Clinch River, Edgemoor and Clinton, Anderson Co.; Clinch River Station, Claiborne Co., Tennessee;
Speers Ferry and Clinchport, Scott Co., Virginia."*^
Powell River, Combs, Claiborne Co., Tennessee.
West of the Mississippi:
Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).
White River, Cotter and Norfolk, Baxter Co., Arkansas (A. A. Hinkley).
Saline River, Benton, Saline Co., Arkansas (H. E. Wheeler).
Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).
Neosho River, near state line, Kansas (R. L. Moodie); Miami, Ottawa Co., Oklahoma (F. B. Isely).
Ten specimens are at hand, but only two or three approach the smooth form, and none is entirely
smooth.
An entirely smooth specimen.
In the upper North Fork of the Holston and the upper Clinch River in Virginia, the var. strigillata gradually replaces the normal form.
56
MEMOIRS OF THE CARNEGIE MUSEUM.
Distribution and Ecology in Pennsylvania (See fig. 6) : This species has been found both in the larger rivers and in some of the smaller creeks, but according to my experience, it is distinctly more frequent in the latter. Yet it does not go up into the extreme headwaters. It is not rare in the Beaver-drainage, and also in French Creek, but entirely absent in the upper Allegheny. At all other localities, it is distinctly a rare shell, and the same is true for the Ohio below our state.
Whenever I found this siiecies alive, it was in swiftly running water, upon bars of gravel, very often in riffies with an abundant growth of Dianthera americana. In fact, on the edge of such patches, I was most successful in taking this species alive. It is mostly not deeply buried, often simply lying upon the bottom. In the Ohio, it is upon the shell-banks, which become accessible only at the lowest stage of the river. This agrees with Scammon (1906), who observed that the ‘‘favorite habitat is bars of gravel or shingle in rather swift current.”
Genercd distribution: Type locality, Wabash River (Say).
Simpson (1900) gives for this species: “Entire Ohio, Cumberland, and Tennes- see river systems; west to Nebraska (doubtful); south to Arkansas and Indian Territory.” The first three rivers undoubtedly are the metropolis of this species, and here it is found from western Pennsylvania'**' through Ohio and West Virginia to Tennessee and northern Alabama (Tennessee-drainage). It is also frequent in the part of Indiana drained by the Ohio, White, Wabash,"*® and Kankakee Rivers (Call, 1896a and 1900), but in Illinois it is found only in the southern part (Ohio and Wabash, Baker, 1906). Thence, in a northerly and northwesterly direction, it is not found any more (this fact should be noted), but it is present farther South
in the Mississippi-drainage, in southern Missouri (Utterback, 1916), in southeastern
*
Kansas, eastern Oklahoma, and in Arkansas. Call (1895, p. 15) reports this species from the Alabama River, Selma, Dallas Co., Alabama. This is the only locality outside the Mississijipi area, and seems to be erroneous, for it has never been found again in this system.
Genus Rotund aria Rafinesque (1820).
Ortmann, 1912, p. 257; Simpson, 1914, p. 903 (as subgenus of Quadrula).
Tyiie Obliquaria tuberculata Rafinesque.
Two species have been assigned to this genus, which may be only varieties of the same species. Only one of them is found in Pennsylvania.
Call (1895) quotes western New York, but this record has never been substantiated. It is missing in Marshall’s list (1895).
According to Goodrich (1914), Q. cylindrica recently has crossed over into the Maumee River, and descended as far as Antwerp, Paulding Co., Ohio. He calls it by the varietal name strigillata, but probably this is an error.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
57
Rotundaria tuberculata (Rafinesquc) (1820).
Quadrula {Rotundaria) tuberculata (Rafinesque) Simpson, 1914, p. 903.
Plate V, fig. 4^
Records from Pennsylvania:
Marshall, 1895 (Allegheny River, Warren Co.).
Rhoads, 1899 (Ohio River, CoraoiDolis, Allegheny Co., and Beaver, Beaver Co.; Beaver River, Wampum,
Lawrence Co.).
Ortmann, 19096, p. 201.
Characters of the shell: Shell rather large and heavy. Outline subrotimd, or ovate, or subqiiadrate, siibtriincate, and sometimes slightly emarginate behind, not very oblique, height and length not very different. Beaks moderately promi- nent, inclined forward. Beak-sculpture consisting of numerous, fine, irregular, broken, or wavy ridges, showing more or less distinctly a zig-zag arrangement, with a posterior triangular loop most distinct, while anteriorly this arrangement is irregular. The first two or three bars are concentric, then follow three or four which are only double-looped, and then other bars (even as many as ten or more), which exhibit the iregiilar zig-zag sculiiture. The beak-sculpture is continued well upon the disc, and is immediately followed by the nodules of the latter. Shell rather compressed, or only slightly swollen, disk gently convex, with an indistinct, posterior ridge, which may be altogether absent. Posterior slope flat or somewhat depressed, often with a more or less distinct wing-like expansion and elevated posterior upper margin (chiefly in young specimens). Surface of shell covered with tubercles or nodules, which, however, always leave free the anterior portion (one-fourth to almost one-half) of the shell. Tubercles very variable in number, size, arrangement, and shape; they may lie rather few, or quite numerous, are always quite irregular in size, and show no definite arrangement. Generally they are a little transverse, at least some of them. In the center of the shell the tubercles are generally most numerous, and upon the posterior slope and the wing, they often assume the shape of radiating, nodulous ribs. In old shells the tubercles disappear toward the lower margin, and in very large specimens the whole lower half of the shell (or even more) may be without tubercles.
Epidermis brown, lighter or darker, uniform in color, only in very rare cases mere traces of broad greenish rays are barely indicated. Growth-rests slightly darker, or not marked.
Hinge-teeth strongly developed and very heavy. Pseudocardinals large, ragged, divergent. Very often there are one or even two subsidiary pseudo- cardinals in the right valve (one in front, the other behind the normal tooth).
58
MEMOIRS OF THE CARNEGIE MUSEUM.
Interclentum well-developed and extremely wide. Lateral teeth short and strong. Beak-cavity very deep, compressed. Dorsal muscle-scars on the hinge-plate. Nacre of a peculiar and characteristic brownish purple color, lighter or darker, sometimes shading to whitish toward the beak-cavity, and coppery-brownish, iridescent posteriorly.
Sexes absolutely indistinguishable in the shell.
L. H. D.
Size: 1. Godfrey, Cat. No. 61.4411 108 mm. 96 mm. 47 mm.
2. Neville Island, Cat. No. 61.1637 77 “ 73 “ 41 “
3. Utica, Cat. No. 61.3947 58 “ 55 “ 36 “
Soft parts (See Ortmann, 1912, p. 258, fig. 7). Gravid females have been found subsequently. Only the outer gills are charged, the placentae are sub- cylindrical and white. Glochidia described and figured by Utterback (1916, PL I, fig. 4)., They are unusually large, 0.267 X 0.325 mm. Surber (1912, PL 2, fig. 19) figures the glochidia of the closely allied R. granifera (Lea). These also are remark- able for their size, 0.29 X 0.355 mm., but I have found them in R. granifera from Black River, Arkansas, to be considerably smaller, 0.25 X 0.28 mm.
Breeding season: I found my gravid females on May 22, 1914; May 25, 1915; July 5, 1913; July 7, 1913; July 13, 1913. Utterback’s glochidia were found on August 11.
Re7narks: Although this species resembles in shape and sculpture several other species, chiefly Quadrula pustulosa and Plethobasus cooperianus, it is always easily recognized by the peculiar color of the nacre and the extremely broad inter- dentiim. I have old specimens, which are so much corroded at the beaks, that the whole section bearing tubercles is gone, and only the smooth lower half of the shell remains intact, yet the interior of the shell characterizes them. All Penn- sylvanian specimens are rather flat, but farther down the Ohio occasional specimens are met with which are somewhat more swollen. A specimen from Portsmouth, Ohio, has distinctly more prominent and more incurved beaks, and might fall under R. granifera (Lea) : but the other characters of the latter, as given by Simpson (1900, p. 795, footnote 2) are absent.
Loealities in Pennsylvania represented in the Carnegie Museum:
Ohio River, Cooks Ferry and Industry, Beaver Co.; Neville Island, Allegheny Co.
Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).
Slipperyrock Creek, Wurtemberg, Lawrence Co.
Allegheny River, Godfrey, Johnetta, and Kelly, Armstrong Co.; Walnut Bend, Venango Co.
French Creek, Utica, Venango Co.
Dunkard Creek, Wiley, Greene Co.
Cheat River, Cheat Haven, Fayette Co.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
59
Localities in Pennsylvania, represented in the Philadelphia Academy of Natural Sciences :
Ohio River, Beaver, Beaver Co.; Coraopolis, Allegheny Co. (S. N. Rhoads).
Other localities represented in the Carnegie Museum:
Lake-drainage:
Lake Erie, La Plaisance Baj", Monroe Co., Michigan (C. Goodrich).
Ohio-drainage:
Ohio River, St. Marys, Pleasants Co., West Virginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio.
West Folk River, Lynch Mines, Harrison Co., West Virginia.
Little Kanawha River, Grantsville, Calhoun Co., West Virginia (W. F. Graham).
Elk River, Shelton, Clay Co., West Virginia.
New River, Hinton, Summers Co., West Virginia.
Tennessee-drainage :
Tennessee River, Tuscumbia, Colbert Co., and Florence, Lauderdale Co., Alabama (H. H. Smith). Shoals Creek, Lauderdale Co., Alabama (H. H. Smith).
Flint River, Gurley, Madison Co., Alabama (H. E. Wheeler).
Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).
Tennessee River, Concord and Knoxville, Knox Co., Tennessee.
French Broad River, Boyd Creek, Sevier Co., Tennessee.
Nolichucky River, Chunns Shoals, Hamblen Co., Tennessee.
Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill and Noeton, Grainger Co.; Austin Mill and Church Hill, Hawkins Co., Tennessee.
South Fork Holston River, Pactolus, Sullivan Co., Tennessee.
North Fork Holston, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia; Mendota, Washington Co., Virginia.
Clinch River, Solway, Knox Co.; Edgemoor, Clinton, and Offutt, Anderson Co.; Clinch River Station, Claiborne Co.; Oakman, Grainger Co., Tennessee; Speers Ferry, Scott Co., Virginia.
West of the Mississippi: c-
James River, Galena, Stone Co., Missouri (A. A. Hinkley).
White River, Hollister, Taney Co., Missouri (W. 1. Utterback); Rogers, Benton Co., Arkansas (R. L. Moodie); Cotter, Baxter Co., Arkansas (A. A. Hinkley).
Distribution and Ecology in Pennsylvania (See fig. 7) : In Pennsylvania, this species has a rather wide distribution, being found in all three river-systems. It goes rather high up (in the Allegheny as far up as Warren Co., according to Marshall, 1895). The smallest streams in which I have found it are Slipperyrock, French, and Dunkard Creeks. From the Monongahela proper records are missing, but it ascends to West Virginia (West Fork River). It is nowhere abundant, and is decidedly one of the rarer shells, only few individuals having been found at any one place. This also holds good for the Ohio below Pittsburgh, where this species is by no means abundant.
GO
MEMOIRS OF THE CARNEGIE MUSEUM.
It is hard to say what the ecological preferences are, but the specimens I col- lected alive always came from riffles with rather coarse gravel and a rapid flow of water. In the Ohio proper it inhabits the shell-banks. However, Baker (1898a, p. 36) says that it is found in the Chicago area in the larger lakes and rivers on a muddy bottom.
General distribution: Type locality, Ohio River (Rafinesque).
The range of this species according to Simpson (1900), is “ Mississippi-drainage generally; southern Michigan; San Saba Co., central Texas.” Call (1885) says: '‘New River, Virginia, to Tuscumbia, Alabama; to Iowa; to Michigan.”
The area occupied by this species includes the Tennessee, Cumberland, and Ohio drainages; toward the west and southwest, it apparently becomes scarce. It has not been reported from Kansas (Scammon, 1906), but is jiresent in southern Missouri and northwestern Arkansas.^^ Simpson’s record from Texas is the
■ Rotundaria tiibercidata. + Plethobasus cooperiamis. • Plethobasus cyphyus.
only one in this state, and the list of Singley (1893) does not contain it. Also in a northwesterly direction, the distribution seems to be limited, and barely reaches to Iowa in the Alississippi. Northwards it extends all over Illinois, to southern Cook Co. (Baker, 1898a, 1906) and goes in the Mississippi, Rock, Wisconsin, and St. Croix Rivers, to Wisconsin (Barnes, 1823; Lapham, 1860; Cooper, 1855). In Indiana it has been recorded by Call (1896a) from the Ohio, Wabash, and White
Call (1895) does not mention this species from Arkansas, but, aside from the localities represented in the Carnegie Museum, it occurs also in the Big Buffalo Fork of White River (Meek & Clark, 1912) and W. I. Utterback (1916) reports it from southern Missouri.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
G1
RiverS; and in Ohio by Sterki (1907a) from the Ohio, Little Miami, and Tuscarawas Rivers. In addition, it is found in southern Michigan (Walker, 1898), and in Lake Erie (Barnes, 1823; Walker, 1913). Here we have apparently the only region, where this species leaves the Mississippi-drainage, and it is quite clear, that this was done by the Wabash-Maumee route. On the Pennsylvanian shores of Lake Erie it has not as yet been found.
Genus Plethobasus Simpson (1900).
Ortmann, 1912, p. 259; Simpson, 1914, p. 805 (as section of Pleurohema) .
Type Obliquaria cyq)hya Rafinesque.
Three species are known to belong to this genus, of which only two are found in Pennsylvania. But since the third turns up in the Ohio, not far from the western state-line, I give here a key for all three of them.
Key to the species of Plethobasus.
ai. Shell rounded, ovate, or oblique, without a radial depression running toward the posterior basal margin.
bi. Shell subrotund, or subovate, only slightly oblique. Disk with scattered, irregularly disposed, rounded, or slightly transverse nodules, leaving free the anterior part of the shell, but generally extending more or less upon the posterior slope P. cooperianus.
62. Shell subovate, very strongly oblique. Disk with more or less transverse tubercles, or sub- concentric, more or less interrupted ridges, restricted to the middle of the shell, leaving the
anterior part as well as the posterior slope free of sculpture P. cicatricosus (extralimital).
a 2. Shell obliquely elongated, with a broad and shallow radial depression running toward the post- basal margin. In front of the depression a radiating row of low and broad, often transverse, tubercles; sometimes also a few indistinct tubercles behind the depression P. cyphyus.
Remarks as to nomenclature and synonyjny: Frierson (1911) has published a paper dealing with these forms, but in my opinion he starts from an incorrect assumption. This assumption is, that U . cicatricosus Say (1829) is the same as U. cesopus Green (1827) (which in turn is identical with cyphyus Rafinesque, 1820). The original description of Say mentions two important characters: tubercles and anterior position of beaks (great obliquity), which do not leave the slightest doubt, that the species was correctly understood by subsequent writers {U. cicatricosus Reeve, 1864, PI. 8, fig. 31, and Simpson, 1900).
Reeve figures {CJ. PI. 13, fig. 50) a smaller and less elongated individual. Frierson believes that this is another species. If that were the case, I would be compelled to make three or four additional species out of my (scanty) material of cicatricosus.
There appears to me to be no ground for Frierson’s introduction of the new
62
MEMOIRS OF THE CARNEGIE MUSEUM.
specific name of delectus {1. c., p. 52, PL 2, lower fig.; PI. 3, upper fig.). This is a synonym of cicatricosus.
I think that Frierson makes another mistake in identifying Lea’s U. varicosus (Obs. I, 1834, PI. 11, fig. 20) with his delectus. Lea’s species is very doubtful, and the figure is possibly inaccurate in several resjiects; but chiefly in that the sculpture is entirely different from that of the species of Plethobasus (continuous, concentric, rather regular ridges, not confined to the middle of the shell). I have shown above that Lea’s species is, if anything, an exceptional individual of Fus- conaia suhrotunda (Lea) (See above, p. 9).
Frierson’s U. cicatricoides (p. 53, PI. 2, upper fig.) is also only an extreme variation of cicatricosus, distinguished by less oblique shell, and less anterior beaks. It connects this species with coopcrianus. I have a specimen (St. Marys, West Virginia), which answers well to Frierson’s figure. A specimen from the same locality is more distinctly oblique, and stands actually midway between the two figures on Frierson’s PI. 2, and comes very close to Call’s figure of U. varicosus (1900, PI. 55) except that it is not so much drawn out at the lower posterior end. And further certain individuals of P. coopcrianus, collected by myself, very closely approach in outline the first one, just mentioned, but the sculpture is more dis- tinctly and typically that of coopcrianus.
Finally these intergrading forms also show certain relationships to P. cyphyus, which will be discussed below.
Plethobasus cooperianus (Lea) (1834).
Qiiadrula cooperiana (Lea) Simpson, 1914, p. 852.
Plate V, fig. 5.
Records from Pennsylvania:
Rhoads, 1899 (Ohio River, Beaver, Beaver Co.; and Coraopolis, Allegheny Co.).
Ortmann, 19096, p. 198.
Characters of the shell: Shell moderately large, heavy. Outline subrotund, subovate, or subtriangular, about as long as high, slightly oblique. Beaks moder- ately high, more or less inclined forward, but not at the anterior end of the shell. Beak-sculpture not observed, but probably poorly developed, and not extending to any considerable degree upon the disk, since the latter is always smooth near the beaks before the tubercles begin. Shell moderately swollen, rather evenly convex upon the sides, without a posterior ridge. Posterior slope very slightly compressed, in young specimens sometimes with an indication of a wing. Disk covered with nodes or tubercles, which, however, always leave free the anterior
ORTMANN; monograph of the naiades of PENNSYLVANIA.
G3
part of the shell, and may be scarce or nearly absent upon the posterior slope. Tubercles variable in size, shape, and arrangement. There is no definite arrange- ment, except sometimes a concentric one, parallel to the growth-lines. The shape of the tubercles may be rounded or tear-like, but generally there are at least some, which are peculiarly compressed and transversely elongated (appearing as “pinched up”). The transverse dilatation of the tubercles is not very great. The tubercles may be rather crowded and numerous, or they may be scarce, sometimes only a few are developed, and they often disappear entirely toward the lower margin. Posteriorly, upon the posterior slope, the tubercles may be missing, or a few, or they may be as numerous as in the middle of the shell, or may even assume the shape of nodular, radiating ribs.
Epidermis yellowish brown to rusty or chestnut-brown, rather dark, when old. Very often the posterior slope is lighter than the rest, yellowish or olive-brown. Growth-rests darker. Greenish, indistinct rays are very rarely indicated and only in young specimens.
Hinge-teeth well-developed. Pseudocardinals divergent, ragged. Interden- tum moderately wide. Lateral teeth moderately long, strong. Beak-cavity moderately deep. Dorsal muscle-scars on the hinge-plate. Nacre whitish, often suffused with a delicate pink inside of the mantle-line, most intense toward the hinge-teeth. Very rarely the whole interior is pink.
No sexual difference whatever in the shells. Call’s remarks (1900, p. 485) about the differences of the males and females do not at all hold good.
L. H. D. •
90 mm. 78 mm. 46 mm.
69 “ 61 “ 38 “
69 “ 71 “ 41 “
Size: 1. Industry, Cat. No. 61.3882. . .
2. Shippingport, Cat. No. 61.3881
3. St. Marys, Cat. No. 61.4595. .
Soft parts (See Ortmann, 1912, p. 361). Breeding season: Gravid females and glochidia have not yet been observed. On June 20 to 22, 1911, I obtained a good number (about fifty) specimens in the. Ohio at St. Marys; additional ones were found on July 13, 1911, at Portland. All had the characteristic light orange color of the soft parts. Of about two dozen females none was gravid, and every one had only the outer gills marsupial. The sexual glands were grayish or brown and in many cases distinctly pinkish. Thus it is probable, that the eggs also are pink, as in P. cyphyus.
Remarks: Externally in shape and sculpture this species greatly resembles Quadrula pustulosa (Lea) and Rotundaria tuberculata (Rafinesque). In the tubercles the resemblance to the latter species is so great that there might be a close genetic
64
MEMOIRS OF THE CARNEGIE MUSEUM.
relationship between the two. However, in the beak-sculpture, in the hinge, the color of the nacre, and chiefly in the soft parts, there are important differences, amounting to generic distinctness. The color of the soft parts of P. cooperianus is also rather unique (pale orange, found also in the other species of the genus). Q. pustulosa is, of course, distinguished by the soft parts; but there are also differ- ences in the shell. It is generally smaller, the tubercles are not of the peculiar transverse shape, and the outline of the shell is not so oblique. The color of the epidermis and the presence of broad rays near the beaks also distinguish Q. pustu- losa.
P. cooperianus is very variable in outline. There are specimens, which are almost round, but generally they are drawn out at the lower posterior end, so as to render the outline ovate and oblicpie, and in some cases even subtriangular. The beaks are always inclined forwards, but they never are at the most anterior end of the shell. Very oblique specimens approach P. cicatricosus (Say), and individuals corresponding to cicatricoides of Frierson (see above), may be regarded to a certain degree as intergrades between the two. In sculpture this species also intergrades in the direction of cicatricosus, if the tubercles are more or less restricted to the middle of the shell. However, although there are certain indi- viduals, which are intermediate in the one or the other character, there is no com- plete series of intergrades, and all my specimens may be assigned to the one or the other species.
Specimens from the Cumberland and Tennessee drainages have the sculpture generally better developed than specimens from the Ohio River.
Localities represented in the Carnegie Museum:
Ohio River, Shippingport, Cooks Ferry, and Industry, Beaver Co., Pennsylvania; Clarington, Monroe
Co., Ohio; St. Marys, Pleasants Co., West Virginia; Marietta, Washington Co., Ohio (V. Sterki);
Parkersburg, Wood Co., West Virginia; Portland, Meigs Co., Ohio.
Cumberland River, Cloyds Landing, Cumberland Co., Kentucky (B. Walker donor).
Tennessee River, Knoxville, and Brabsons Ferry, Knox Co., Tennessee.
Clinch River, Edgemoor, Anderson Co., Tennessee.
Distribution and Ecology in Pennsylvania (See fig. 7) : This is one of the rarest species in Pennsylvania, and exists only in the Ohio River in Beaver and Allegheny Cos., and is even there very scarce. I was able to secure only four specimens, one of which was alive. The latter was found upon a shell-bank, in a steady current, associated with the usual bank-forming species.
Farther down the Ohio, P. cooperianus becomes more abundant, and it is cpiite common in the region of St. Marys and Marietta, where it is frequently taken
ORTMANN: monograph of the naiades op PENNSYLVANIA.
65
by the clam-diggers out of deep water, with the other bank-forms. This seems to be its favorite habitat.
General distribution: Type-locality , Ohio River (Lea).
This species seems to be restricted to the Ohio, Ciimlierland, and Tennessee systems, and the known records are all from these rivers and some of their larger tributaries, as for instance the Cdinch in eastern Tennessee, the Wabash in Illinois and Indiana, and possibly also the Illinois River (Fulton Co.) in Illinois (Baker, 1906). It has been reported also from the Mississippi in lowa,^^ but this is doubted by Simpson. It is to be especially noted, that from Indiana through Ohio and West Virginia to Pennsylvania, this species is restricted to the Ohio proper.
Plethobasus cyphyus (Rafinesque) (1820).
Pleurobenia cesopus (Green) *Simpson, 1914, p. 806; Pleurohema cyphia (Rafin- esque) Vanatta, 1915, p. 556.
Plate V, fig. 6; Plate VI, figs. 1, 2, 3.
Records from Pennsylvania:
Green, 1827 (rivers near Pittsburgh).
Harn, 1891 (western Pennsylvania).
Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co.; and Beaver River, Wampum, Lawrence Co.). Ortmann, 19096, p. 198.
Characters of shell: Shell rather large, heavy. Outline elongated-subovate, generally (except in young shells) distinctly longer than high, somewhat oblique, with the beaks not very high, inclined forward, but not at the anterior end of the shell. Beak-sculpture consisting of a few, thick, concentric ridges, not extending upon the disk. Shell moderately swollen, but old shells sometimes more con- siderably so; sides convex, with a broad and shallow radial depression, running from the beaks toward the posterior lower margin, producing a more or less distinct shallow excavation of the margin. In front of this depression, a radial row of low, transverse tubercles running from the beaks to the lower margin. These tubercles begin at a certain distance from the beaks, are more or less distinct (sometimes almost obsolete), and are somewhat irregular, larger or smaller. In rare cases they are cut by obscure, nearly vertical, lines (similar to P. cicatricosus) . Behind the radial depression there is a low and broad ridge, passing gradually into the posterior slope. This part of the shell is generally less swollen than the anterior, and may be entirely smooth, or may show irregular, low rugosities, and sometimes
Call (1885) and Keyes (1888) give Muscatine, Iowa, and Pratt (1876) Davenport, Iowa, but it is not in Witter’s (1878) list from Muscatine,
66
MEMOIRS OF THE CARNEGIE MUSEUM.
there are individuals with indications of another row of tubercles upon the posterior ridge, which is, however, always less distinct than the anterior row.
Epidermis yellow to dark brown. In young specimens, the color is- generally a rather light yellow (wax-yellow), becoming russet or chestnut-brown in older ones. There is no trace of rays. Growth-rests darker than the rest of the surface, but not sharply marked.
Hinge-teeth well-developed. Pseudocardinals moderately large, divergent, ragged. Interdentum moderately wide or narrow. Lateral teeth rather long. Beak-cavity not very deep. Dorsal muscle-scars on the hinge-plate close to the beak-cavity. Nacre white.
There is not the slightest sexual difference in the shell, and neither the length
nor the obesity is connected with the sex.
L. H. D.
Size: 1. Kelly, Cat. No. ()1.3817 122 mm. 81 mm. 50 mm.
2. Kelly, Cat. No. 01.4598 (long cf) 108 “ 74 “ 47 “
3. Kelh", Cat. No. 61.5560 (long 9, gravid) 93 “ 64 “ 43 “
4. Kelly, Ctit. No. 61.3817 (short 9 , gravid) 90 “ 07 “ 42 “
5. Godfre„v, Cat. No. 61.4599 (short d") 88 “ 72 “ 45 "
6. Shippingport, Cat. No. 61.4600 (long cf) 78 “ 58 “ 36 “
iiojt parts (See Ortmann, 1912, p. 260, fig. 8). Glochidia (See Ortmann, ibid.). It should be remarked that the measurements previously given were taken from immature glochidia. Gravid females collected subsequently contained fully developel larva?, and according to these, the glochidia measure 0.21 X 0.20 mm. Surber (1912, PI. 1, fig. 12) illustrates them, and gives the size as 0.22 X 0.20 mm.
Breeding season: I have found gravid females on the following dates: June 20,1911; June 21, 1911; June 22, 1909; July 3, 1908; July 4, 1911; July 13, 1908; July 13, 1911; July 25, 1910; July 27, 1910. Glochidia were observed on July 4 and July 25. Thus this species appears to be typically tachytictic, breeding in June and Jul}^ Probably the season begins in May, as stated by Surber.
Remarks: This is a well-marked species, at least in Pennsylvania, character- ized by the shape of the shell, sculpture, and color of the epidermis and soft parts. The most closely allied species is P. cicatricosus, a form not found in Pennsylvania, but farther down the Ohio (nearest point St. Marys, West Virginia). P. cyphyus is distinguished from this species by the more elongated, -less oblique shape, with the beaks less anterior, by the sculpture, and the presence of an oblique furrow upon the disk. However, there are individuals, which form, to a degree, a transi- tion between these two species. To these transitional forms belongs U. cicatricoides of Frierson (See above, p. 62), but, on account of the absence of a radiating furrow.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
67
the latter should be classed rather with P. cicatricosus. One of my specimens from St. Marys agrees rather well with U. cicatricoides, but it has upon the disk a slight indication of a depression, since the tubercles are cut off rather suddenly behind, making the posterior part of the shell appear contracted; but the liroad furrow of P. cyphyus is not by any means developed.
In sculpture, P. cyphyus represents a further reduction of the tubercles of P. cicatricosus. They are rather poorly developed, but still show in most cases the character of short, transverse ridges. Sometimes these tubercles have almost entirely disappeared. Where they are better developed, in rare cases even the vertical dividing lines of the ridges, seen so frequently in P. cicatricosus, are indi- cated.
Young specimens of P. cyphyus somewhat resemble in shape Truncilla rangiana (Lea). But the latter species has a more greenish olive epidermis, with more or less distinct rays.
There is considerable variation in the comparative length of the shell of P. cyphyus. A short appearance is given to the shell by a smaller development of the posterior slope, and such shells are decidedly more abundant among young individuals. Old specimens are, as a rule, greatly elongated, and in these it often happens that the radial furrow becomes effaced. If such specimens have at the same time poorly developed tubercles, they resemble in shape old specimens of Fusconaia subrotunda, but they generally differ in the color of the epidermis, lighter hinge, and chiefly in the color of the soft parts (light orange). Of course, if females are at hand, no mistake is possible.
Localities in Pennsylvania, represented in the Carnegie Museum:
Ohio River, Shipping-port and Industry, Beaver Co.; Coraopolis, Allegheny Co. (S. N. Rhoads). Monongahela River, Westmoreland Co. (G. A. Ehrmann).
Allegheny River, Aladdin, Godfrey, Kelly, and Templeton, Armstrong Co.
Locality in Pennsylvania represented in the Philadelphia Academy of Natural Sciences:
Beaver River, Wampum, La-wrence Co. (S. N. Rhoads).
Other localities represented in the Carnegie Aluseum:
Ohio-drainage:
Ohio River, Toronto, Jefferson Co., Ohio; St. Marys, Pleasants Co., West Virginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio.
Tuscara-was River, Ohio (Holland collection).
Wabash River, New Harmony, Posey Co., Indiana (A, A, Hinkley),
68
MEMOIRS OF THE CARNEGIE MUSEUM.
Te7inessee-drainage:
Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).
French Broad River, Boyd Creek, Sevier Co., Tennessee.
Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Holston Station, Grainger Co., Tennessee.
North Fork Holston River, Rotherwood, Hawkins Co., Tennessee.
Clinch River, Solway, Knox Co.; Edgemoor, Clinton, and Offutt, Anderson Co.; Black Fox Ford, Union Co.; Clinch River Station, Claiborne Co.; Oakman, Grainger Co., Tennessee; Clinchport, Scott Co., Virginia.
Powell River, Combs, Claiborne Co., Tennessee.
West of Mississippi:
Meramec River, Meramec Highlands, St. Louis Co., ^Missouri (N. M. Grier).
Distribution and Ecology in Pennsylvania (See fig. 7) ; This species is found in Pennsylvania, preeminently in the Ohio and Allegheny Rivers; from the localities in the Monongahela and Beaver, only single individuals are known. It prefers riffles with coarse gravel and strong current, and in Beaver Co. it is found upon the shell-banks in a strong and steady current. The same is true farther down the Ohio, where it is often obtained by the clam-diggers.
General distribution: Type locality, Falls of the Ohio (Rafinesque) at Louis- ville, Kentucky.
In the Allegheny^ River, at Templeton, Armstrong Co., and in the Mononga- hela, in Washington and Westmoreland Cos., Pennsylvania, this species reaches its highest points of advance in an upstream direction. Also Beaver River at Wampum, Ijawrence Co., is an extreme point. Farther westward, its range follows the Ohio, and embraces, according to Simpson (1900), the Ohio, Cumber- land, and Tennessee river-systems; west to Missouri and hlinnesota. In the upper Ohio, it goes very little into the tributaries, and is known only from the Tuscarawas and Scioto in Ohio (Sterki, 1907a^®), Wabash (Call, 1896a and 1900), and White River (Lea, Obs. X, 1863, p. 432) in Indiana. In Illinois it covers a large territory (Illinois River up to Kankakee and Fox Rivers). It also follows up the Mississippi and its tributaries in northern Illinois and eastern Iowa (Des IMoines River, Witter, 1878, Call, 1885), and goes as far as Minnesota (Grant, 1886, and Holzinger, 1888).
West of the Mississippi, records are scarce or doubtful,®^ but the Carnegie iMuseum has it from the jMeramec River, near St. Louis, and Utterback (1916)
Sterki gives also the Mahoning River. I question this record, since I have never seen a trace of it in the lower Mahoning in Pennsjdvania, and since Dean (1890) does not mention it.
Scammon gives it as rare from Verdigris River in Kansas. A specimen from Wakarusa River, Lawrence, Douglas Co., Kansas, determined by Scammon as P. cesopiis, and sent to the Carnegie Museum by R. L. Moodie, agrees very well with Scammon’s figure (1908, pi. 78, fig. 2) but it is not this si^ecies. Our specimen is an old, dead shell, and probably an abnormal Quadrula quadrula.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
69
mentions at least one place in western Missouri (Little Blue River, Courtney, Jackson Co., near Kansas City). The species has never been reported from the region of the lower Mississippi, and although Call (1896a and 1900) includes the Alabama River in its range, this record has not been confirmed by anybody else (See Lewis, 1877).
Genus Pleurobema Rafinesque (1820).
Ortmann, 1912, j). 261; Simpson, 1914, p. 732.
Type Unio clava Lamarck.
The conception of this genus has been changed greatly in recent times. In Pennsylvania, I distinguish two species, but one of them is quite polymorphous, with a number (five) of variations and ecological (or local) races or varieties.
Key to the Species and Varieties of PlexTrobema. tti. Shell rather large, upright, or oblique. Epidermis chestnut to dark brown or blackish, with in- distinct capillary rays, which generally are not broken up into blotches. bi. Shell not very oblique, more or less upright. Nacre white or red. Radial furrow strong, weak, or absent. Diameter variable.
Cl. Furrow strong or weak, but always present. Nacre white, very rarely red. Diameter mostly over fifty percent of length (very rarely less). Shape upright.
di. Outline subtriangular; furrow strong P. obliquuni.
di. Outline rounded; furrow generally weak, but present . . .P. obliquum cordatum.
C2. Furrow generally absent (rarely a trace seen). Nacre white or red. Diameter variable. Shape subtriangular, upright, or slightly oblique. di. Shell rather large. Growth lines irregular and indistinct. Epidermis dark.
Cl. Nacre white, rarely red. Diameter fifty percent of length or more.
P. obliquum catillus.
Ci. Nacre white, salmon, or red. Diameter less than fifty percent of length.
P. obliquuvi coccineuni.
do. Shell smaller. Growth lines more distinct and regular. Epidermis lighter.
P. obliquum pauperculum.
&2- Shell very oblique, beaks directed anteriorly. Nacre generally red, rarely white. Furrow strong,
rarely weak. Diameter mostly over fifty percent of length P. obliquum rubrum.
Ui. Shell rather small, very oblique, beaks located well anteriorly. Epidermis j^ellowish or light brown, with distinct rays, which are often broken up into blotches P. clava.
Pleurobema obliquum (Lamarck) (1819).
Quadrula obliqua (Lamarck) Simpson, 1914, p. 881.
Plate VI, figs. 4, 5, 8.
Records from Pennsylvania:
Call, 1885 (Allegheny River).
Harn, 1891 (western Pennsylvania).
70
MEMOIRS OF THE CARNEGIE MUSEUM.
Clapp, 1895 (Allegheny Co.).
Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co.).^^
Ortmann, 19096, p. 199.
Characters of the shell: Shell rather large, heavy. Outline subtriangular, rounded before, subangular and more or less drawn out at the lower posterior end, with the lower margin gently convex in the anterior part, and generally concave in the posterior part, and with the supero-posterior margin gently curved. Beak more or less elevated, and swollen, directed obliquely forward, or more or less incurved, situated nearer to the anterior end of the shell. Beak-sculpture rudi- mentary, consisting of two or three indistinct, concentric bars. Shell rather swollen anteriorly, the diameter generally over fifty percent of the length. Posteriorly the shell is compressed, with a broad, more or less distinct, radial furrow, running from the beaks toward the lower posterior margin, producing there a shallow emargination. Surface of shell smooth, without any sculpture. Behind the radial depression there is a rounded, low elevation, which, however, does not form a dis- tinct ridge, and passes insensibly into the posterior slope.
Ejiidermis lighter or darker brown, generally of a dark chestnut hue, blackish, when old, with fine and indistinct green or blackish capillary rays, or bundles of rays, which are mostly entirely obliterated in old specimens. The rays do not break up into blotches. Growth-rests more or less distinct, darker than the rest of the shell.
Hinge-teeth well-developed, rather strong. Pseudo cardinals divergent, ragged. Interdentum rather wide. Lateral teeth moderately long. Beak-cavity more or less deep, often somewhat compressed. Dorsal muscle-scars on the hinge-plate. Nacre mostly white, rarely reddish.
No sexual differences in the shell.
|
L. |
H. |
D. |
Pr.ct. |
|
|
1. Industry, Cat. No. 61.3900 (unusually |
||||
|
produced) |
. . . 109 mm. |
82 mm. |
46 mm. |
.42 |
|
2. Industry, Cat. No. 61.3902 |
. . . 90 “ |
77 “ |
.51 “ |
.57 |
|
3. Godfrey, Cat. No. 61.3901a |
. . . 83 “ |
69 “ |
42 “ |
.51 |
|
4. Charleroi, Cat. No. 61.2869 |
. . . 59 “ |
52 “ |
33 “ |
.56 |
Soft parts (See Ortmann, 1912, p. 264). I havn examined nearly a thousand individuals of this species, and have found the structure of the soft parts as de- scribed, with only the outer gills marsupial. However, a few exceptions have been noted in the case of five specimens received through the courtesy of Dr. R.
As has been stated by the writer (19096, pp. 186 and 199) only part of the specimens from Cora- opolis, recorded by Rhoads, belong to PI. ohliquum, the rest are Fusconaia subrotunda, while all of Rhoads’ specimens from Beaver (in the Philadelphia AcademjO are F. subrotunda.
ORTMANN: monograph of the naiades of PENNSYLVANIA.
71
E. Coker, by whom they were collected in the Cumberland River, near Clarksville, Montgomery Co., Tennessee. In the extreme case about half of the inner gills, in their middle portion near the margins, was marsupial; in three other cases, much smaller sections of the inner gills were charged, and in one case a few eggs were near the central margin of the right inner gill. Along with these abnormal specimens, two were received with normal marsupium, but none had all four giWs fully charged, as is the case in the genera Fusconaia, Quadrula etc.®^ It should be borne in mind that these specimens were selected and sent to me to demonstrate the fact that sometimes there are eggs in the inner gills. This observation does not invalidate the rule that this species normally has only the outer gills marsupial. Since the genus Pleurobema undoubtedly is descended from ancestral forms, which had all four gills marsupial (as for instance Fusconaia)^ it is not astonishing, that the present species sometimes reverts, although not fully, to this old condition {Atavism). From the genus Fusconaia, to which it is most closely allied by the shell, it differs nevertheless by the fact that the placentae are always lanceolate, not subcylindrical, and that their color is always white, never red.
Glochidia (observed by myself only in the specimens just mentioned) agreeing in shape and size with those of P. ohliqumn coccineum: 0.15 X 0.15 mm. Ac- cording to Surber (1915) they measure: 0.160 X 0.175 mm.
Breeding season: I have gravid females collected on the following dates: June 3, 1911; June 20, 1911; June 21, 1911; June 24, 1909. On the first date a specimen with glochidia was obtained. The species is apparently tachytictic, but the duration of the breeding season is not fully known. Surber’s specimens with glochidia were collected on July 14.
Remarks: A quite characteristic species, in its typical phase (called pig- toe” by the clam-diggers), but subject to an immense range of variation. The variations concern chiefly the general shape. While the typical form (See PI. VI, fig. 4) is characterized by its subtriangular outline, by the rather upright shape, with the gently incurved beaks not situated at the anterior end, and by the well- developed radial furrow, all these characters may vary. The commonest vari- ation is that in which the posterior lower angle becomes more or less produced, chiefly occurring in old shells (PL VI, fig. 8) so that the diameter becomes unusually low. It is, however, observable that such specimens, when young, were quite normal. In other specimens the beaks are more upright, making the whole shell appear higher and shorter. The radial depression is extremely variable, often different on the two sides of the same specimen, and tends in many specimens to
Lefevre & Curtis (1912, p. 120) enumerate this species under the “ Tetragente,” where the mar- supium comprises all four gills.
72
MEMOIRS OF THE CARNEGIE MUSEUM.
become effaced, leading thus to the var. catillus (PL VI, figs. 5, 6). Again the posterior angle of the shell may be less developed, and the posterior end may be shorter, a condition generally connected with a deficient development of the whole posterior slope, thus emphasizing the swollen anterior part of the shell. In such cases, the lower margin is more curved upward behind, producing a more rounded, not triangular outline, leading to the var. cordatum. The obesity of the shell, although always considerable, is variable, but there are no shells which could be called flat; yet in specimens with the posterior angle much drawn out, the posterior section of the shell may be flat (PL VI, fig. 8). The nacre is generally white, but individuals are occasionally found, where it is reddish.
Although the group of P. obliquum has been misunderstood by most authors, the main species has generally been properly recognized, and the references given by Simpson are correct. But the figure of Call (1900, PL 59, upp. fig.), under the name of U. solidus, should be added: it is a typical P. obliquum.
• Pleurobema obliquum.
+ Pleurobema obliquum cordatum.
■ Pleurobema obliqimm catillus.
X Pleurobema obliquum coccineum.
V Pleurobema obliquum paupercuhun.
A Pleurobema obliquum rubrum.
Localities in Pennsylvania represented in the Carnegie Museum:
Ohio River, Smiths Ferry (W. F. Graham), Shippingport, Cooks Ferry and Industry, Beaver Co.; Corao- polis (S. N. Rhoads) and Neville Island, Allegheny Co.
Monongahela River, Westmoreland Co., and Charleroi, Washington Co. (G. A. Ehrmann).
Allegheny River, Godfrey and Kelly, Armstrong Co.
ORTMANN: monograph op the naiades of PENNSYLVANIA.
73
Other localities represented in the Carnegie Museum:
Ohio-drainage :
Ohio River, Beech Bottom, Brooke Co., West Virginia (W. F. Graham); Toronto, Jefferson Co., Ohio; Clarington, Monroe Co., Ohio; St. Marys, Pleasants Co., West Virginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio.
Tuscarawas River, Ohio (Holland collection).
Cumberland- Tennessee-drainage :
Cumberland River, Clarksville, Montgomery Co., Tennessee (R. E. Coker donor).
Tennessee River, Tuscumbia, Colbert Co., and Florence, Lauderdale Co., Alabama (H. H. Smith). Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).
Tennessee River, Knoxville and Brabsons Ferry, Knox Co., Tennessee.
French Broad River, Boyd Creek, Sevier Co., Tennessee.
Holston River, Mascot, Knox Co.; Hodges, Jefferson Co.; Noeton and Holston Station, Grainger Co., Tennessee.
Clinch River, Solway, Knox Co.; Edgemoor, Clinton, and Offutt, Anderson Co., Tennessee; Needham Ford, Union Co., Tennessee (B. Walker donor).
Distribution and Ecology in Pennsylvania (See fig. 8) : The typical form of P. obliquum is restricted in Pennsylvania to the three large rivers, the Ohio, Alle- gheny, and Monongahela. In the Allegheny it is found as high up as Armstrong Co., but is not very abundant there. In the Monongahela, judging from the large number of specimens collected by G. A. Ehrmann, it must have been at one time abundant in the vicinity of Charleroi, but we do not know how far it ascended the river. In the Ohio below Pittsburgh it is a common shell. Farther down, it is extremely abundant, and is the prevailing species, at leatet locally. It is the shell, which largely contributes in forming the shell-banks, in rather deep, steadily flowing water, and, next to the “mucket” (Actinonaias ligamentina) , it is the shell most highly valued by the clam-diggers. In Pennsylvania, it is present also in riffles, and immediately above them, in strong current, and among coarse gravel.
General distribution: Type locality, Ohio River (Lamarck).
On account of the great confusion prevailing with regard to this species, it is hard to make out the limits of its distribution, but it is certain that it chiefly inhabits the systems of the Ohio, Cumberland, and Tennessee Rivers. In the Ohio-drainage it does not go much into the tributaries, but is known from the Muskingum-Tuscarawas River in Ohio (Sterki, 1907a), and from the Wabash in Indiana (Call, 1896a and 1900). It occurs also in the Mississippi in Illinois (Baker, 1906), and goes northward into Wisconsin and Minnesota. It also has been reported from the Alabama River (Lewis, 1877, and Simpson, 1900), but this is doubtful. I have not seen any specimens of it in the collections made there by H. H. Smith.
74
MEMOIRS OF THE CARNEGIE MUSEUM.
The existence of typical P. obliquum becomes uncertain in the west and south- west. It has not been recorded by Scammon (1906) from Kansas, and although Call (1895, p. 32) cites it from the Ouachita River in Arkansas, I have never seen a genuine P. obliqumn in all the rich material I have received from H. E. Wheeler taken from this river. In this region it seems to be entirely replaced by the vars. catillus and rubrurn. Utterback (1916) failed to find this species in Missouri.
From what we know at present it is quite evident that the species prefers the larger rivers (See Call, 1900, p. 502) and thus it is easily understood why it has not migrated from the Mississippi-Ohio system into any other drainage.
Pleurobema obliquum cordatum (Rafinesque) (1820).
Qaadrula plena (Lea) Simpson, 1914, p. 886; Quadrula cordata (Rafinesque)
Vanatta, 1915, p. 558.
Plate VII, fig. 1.
No records from Pennsylvania previous to those given below.
Characters of variety: Much like typical obliquum, but shape more rounded, which is due to the slight development of the lower posterior angle and the whole posterior slope. In consequence of this, the shell appears more elevated and more upright. With regard to the diameter, the shell resembles P. obliquum (over fifty percent of length). The radial furrow is generally weak, but present. Nacre white, or slightly pink.
|
L. |
H. |
D. |
Pr.ct. |
||
|
Size: 1. Cooks Fe |
iiTv, Cat. No. 61.44346 |
78 mm. |
79 mm. |
51.5 mm. |
.66 |
|
2. Godfrey, |
Cat. No. 61.39016 |
70 “ |
67 “ |
43 |
.61 |
|
3. Godfrey, |
Cat. No. 61.4378 |
56 “ |
57 “ |
34 |
.61 |
Soft parts and glochidia not observed in Pennsylvania. In the region of the upper Tennessee I found a few gravid females, but none with glochidia. The anatomy is the same as in the main species. Only the outer gills possess marsupial structure.
Breeding season: Gravid females were found on May 25, 1914.
Remarks: This form hardly deserves a varietal name, and I retain it only because it has been distinguished as a “species” by previous authors. In Penn- sylvania, it is nothing but an individual variation, found very rarely, and always associated and intergrading with the typical form. The three specimens, of which measurements are given, are the only ones I have from Pennsylvania, and even these are not quite typical representatives of cordatum. In the upper Tennessee region, I have found this form more frequently, but also there it is by no means a well-defined race, and I know of no region, where cordaturn is found pure.
ORTMANN; monograph op the naiades of PENNSYLVANIA.
75
Localities represented in the Carnegie Museum:
Ohio-drainage:
Allegheny River, Godfrey, Armstrong Co., Pennsylvania.
Ohio River, Cooks Ferry, Beaver Co., Pennsylvania.
Tennessee-drainage :
Tenne,ssee River, Florence, Lauderdale Co., Alabama (H. H. Smith); Bridgeport, Jackson Co., Alabama (B. Walker, donor).
French Broad River, Boyd Creek, Sevier Co., Tennessee.
Holston River, Mascot, Knox Co.; Hodges, Jefferson Co.; ITolston Station, Grainger Co., Tennessee. Clinch River, Solway, Knox Co.; Edgemoor and Clinton, Anderson Co., Tennessee.
Distribution (See fig. 8) : Type locality, Ohio River (Rafinesque) .
Wherever found, in Pennsylvania, as well as in the Tennessee-drainage, this form is associated with the main form, and does not show any remarkable facts in its distribution.
Pleurobema obliquum catillus (Conrad j (1836).
Quadrula solida (Lea) Simpson, 1914, p. 885; Pleurobema obliquum catillus (Con- rad) and Pleurobema catillus (Conrad) Utterback, 1916, p. 79, 82.
Plate VI, fig. 6; Plate VII, fig. 2.
No records from Pennsylvania previous to those given below.
Characters of variety: This form may be briefly characterized as a P. obliquum, in which the radial furrow is obliterated. In all other respects it resembles the main species, and varies in the same way. The shell is more or less subtriangular, upright or somewhat oblique. The diameter is always considerable, fifty percent or more of the length. The nacre is generally white, rarely pinkish.
|
L. |
H. |
D. |
Pr.ct. |
||
|
Size: 1. |
Industry, Cat. No. 61.3895 |
83 mm. |
65 mm. |
48 mm. |
.58 |
|
2. |
Industrv, Cat. No. 61.3895 |
81 “ |
69 “ |
45 “ |
.56 |
|
3. |
Godfrey, Cat. No. 61.4567 |
61.5" |
51 ■" |
32 “ |
.52 |
Soft parts: Identical with those of the main form, and also the glochidia.
Breeding season: Gravid females were found on May 22, 1914; June 22, 1909; June 24, 1909. Glochidia on June 24. Specimens from the White River in Ar- kansas were gravid, containing eggs on August 2 and 5, 1914.
Remarks: This form likewise is not sharply separated from the main species, and is connected with it by very gradual transitions, but it has a better claim to be recognized as a variety than var. cordatum, since it seems to replace the main species at least in certain regions. It has been largely misunderstood by previous authors.
76
MEMOIRS OF THE CARNEGIE MUSEUM.
and even Simpson did not recognize it clearly. In fact, two of the specimens of which he published the measurements (the first two) fall under the var. coccineum. The s^monymy given by Simpson is correct, as far as it goes. But there is no doubt that U. catillus Conrad (1836), which is made by Simpson a synonym of U. coc- cineus should be united with U. solidus Lea. Its diameter is fifty-one percent of the length (according to Conrad’s figure, PL 13, fig. 2), and its greater obesity is precisely the character which distinguishes it from U. coccineus. U. solidus (ac- cording to Lea’s figure) has a diameter of sixty percent of the length, while U. coccineus Conrad {ibid., PI. 13, fig. 1) has the diameter thirty-seven percent of the length.
That U. catillus more nearly approaches U. solidus, was already recognized by Utterback, who also clearly saw that these forms in Missouri pass into each other and into obliquum. Utterback tried to express this in a very peculiar way l\y naming one of the intergrades P. catillus, and another P. obliquum catillus, but this can hardly find approval.
The Carnegie Museum ])ossesses, from the Hartman collection, a specimen (Cat. No. 61.1440) from Cincinnati, labeled U. solidus, “type” (meaning typical), and “Lea datum.” This, consequently, is an authentic specimen from Lea, from one of the type localities. It has a diameter of sixty-four percent, and thus cor- responds closely with Lea’s original figure, and with my conception of this form.
That U. solidus of Call (1900, PI. 59, upp. fig.) is a typical P. obliquum, has been mentioned above. What Scammon (1906, PI. 85) figures as a young Quad- rula solida, is not this, but a rather good P. obliquum rubrum.
Unio fulgidus Lea (Obs. IV, 1848, PL 4, fig. 10), supposedly from Alexandria, Louisiana, is made by Simpson a synonym of Q. solida. It is founded upon a very young specimen, and I hardly think that it is possible to make out what it really is. However, Walker has communicated under the name of fulgidus a specimen from Iowa City, which undoubtedly belongs to solidus (= catillus). It has a rather shining epidermis, but otherwise closely resembles specimens labeled solidus by Walker. Such specimens seem to prevail west of the Mississippi, but I have them also from the Tennessee-drainage.
In Pennsylvania, P. obliquum catillus may be recognized by the absence of the radial furrow, the sutriangular outline, and considerable obesity. By the reduc- tion of the latter, i.e., when the shell becomes more compressed, it passes very gradually into the var. coccineum, and it