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.

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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.

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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.

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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

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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,

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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.

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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 is possible to separate the transitional
forms only by exact measurements, and an artificial dividing line. I have drawn
this line at the diameter of fifty percent of the length, but I again must emphasize

ORTMANN; monograph of the naiades of PENNSYLVANIA.

77

that this is arbitrar}^, separating things which are actually connected. However,
for practical purposes (naming of specimens) it is a great help, and serves to simplify
matters.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Shipping-port, Cooks Feny, and Industry, Beaver Co.; Coraopolis, Allegheny Co. (S. N.
Rhoads).

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Mahoning River, Mahoningtown and Coverts, Lawrence Co.

Allegheny River, Natrona, Allegheny Co.; Aladdin, Godfrey, Johnetta, Kelly, and Templeton, Arm-
strong Co.; Walnut Bend, Venango Co.; Hickory, Forest Co.; Warren, Warren Co.

Cheat River, Cheat Haven, Fayette Co.

Other localities represented in the Carnegie Museum:

Ohio-drainage :

Ohio River, Toronto, Jefferson Co., Ohio; Steubenville, Jefferson Co., Ohio (W. F. Graham); St. Marys,
Pleasants Co., West Virginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio; Cincin-
nati, Hamilton Co., Ohio (Hartman collection).

Tuscarawas River, Ohio (Holland collection).

Tennessee-drainage :

Duck River, Columbia, Maury Co., Tennessee (B. Walker, donor).

Tennessee River, Florence, Lauderdale Co. (H. H. Smith); Bridgeport, Jackson Co., Alabama (B.
Walker, donor).

Holston River, Noeton, Grainger Co., Tennessee.

Clinch River, Edgemoor, Anderson Co.; Black Fox Ford, Union Co., Tennessee.

West of Mississippi:

Iowa River, Iowa City, Johnson Co., Iowa (B. Walker, donor).

Meramec River, Mer'amec Highlands, St. Louis Co., Missouri (N. M. Grier).

Marais des Cygnes River, Rich Hill, Bates Co., Missouri (W. I. Utterback).

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

White River, Cotter and Norfolk, Baxter Co., Arkansas (A. A. Hinkley).

Black River (H. E. Wheeler) and Spring River (A. A. Hinkley), Black Rock, Lawrence Co., Arkansas.

From other localities in southern Arkansas (Ouachita River) and in Oklahoma, similar forms are
at hand, but I omit them here, since they require further study, chiefly with regard to their relation to
the southwestern forms of P. obliquum rubrum, which are not at all clear. (See Wheeler, 1918, p. 124.)

Distribution and Ecology in Pennsylvania (See fig. 8) : In the Ohio and Alle-
gheny this form is found associated with typical P. obliquum. No specimens
have come to hand from the Monongahela proper, but it must have existed there,
since it turns up in the lower Cheat. In the Allegheny, it goes a great distance
beyond the range of P. obliquum, up to Warren Co., but in this whole region it is
associated with, and passes into, P. obliquum coccineum. It also has entered the
Beaver River, and goes even up into the lower part of the Mahoning, passing again

78

MEMOIRS OF THE CARNEGIE MUSEUM.

into coccinewm. It is interesting to see that it actually exists in the . Mahoning,
for one of the original localities mentioned by Lea for his U. solidus, is “Mahoning
River, Ohio.”

In the lower Allegheny and in the Ohio just below Pittsburgh, this form is
even more abundant than the normal P. obliquum. It is found under the same
conditions as the latter, but most abundantly upon gravel bars with strong current.

General distribution: Type locality, Scioto River, Ohio (Conrad).

This variety seems to be found wherever the typical form is present, that is to
say, in the Ohio-, Cumberland-, and Tennessee-drainages; but particulars cannot
be accurately expressed on account of the confusion which prevails with regard
to it. It is certainly present in the upper Tennessee-drainage, going upstream
about as far as P. obliquum. In addition this variety seems to be rather frequent
west of the Mississippi, in regions where the typical P. obliquum is largely absent
or doubtful, namely in Iowa, Missouri, and northern Arkansas. As has been
indicated above, P. catillus seems there to replace P. obliquuyn, and in turn connects
with the var. rubrum. This group of forms {catillus-rubrum) develops farther to
the southwest into a peculiar assemblage, which cannot be discussed here. In
Missouri there is shown a tendency on the part of catillus to pass into coccineum
in small streams, exactly as is the case in Pennsylvania.

Pleurobema obliquum coccineum (Conrad) (1836).

Quadrula coccinea (Conrad) Simpson, 1914, p. 883.

Plate VII, figs. 3, 4, 5.

Records from Pennsylvania:

Conrad, 1836 (Mahoning River, near Pittsburgh). “

Marshall, 1895 (Allegheny River, Warren Co.).

Rhoads, 1899 (Beaver River, Wampum, Lawrence Co.).

Ortmann, 19096, p. 200.

Characters of variety: Differs from P. obliquum by the flat and compressed
shell, less prominent beaks, and the absence of the radial furrow; In the latter
character, it agrees with P. obliquum catillus, but differs from this in the com-
jiression, the diameter being less than fifty percent of the length. The shape of
the shell is subtriangular or, on account of the low beaks, more or less subovate.
Color of nacre very variable: white, salmon, or all shades of pink to deep red.

The nearest point of the Mahoning to Pittsburgh is where it joins the Shenango to form the
Beaver, at Mahoningtown, Lawrence Co., Pa. I have found this form at this place, and this should be
regarded as the Type locality.

I

ORTMANN: monograph of the naiades of PENNSYLVANIA. 79

L.

11.

D.

Pr.ct.

1. Mt. Morris, Cat. No. G1.4574

Ill ntm.

88 mm.

43 mm.

.39

2.’ Larabee, Cat. No. 61.3350

100 “

81 “

49 “

.49

3. Eastbrook, Cat. No. 61.3349

88 “

67 “

32 “

.36

4. Mahoningtown, Cat. No. 61.3907. . . .

69 “

59 “

32 “

.46

Soft parts (See Ortmann, 1912, p. 263) : I have seen several hundred females,
many of them gravid, but in no case I have found marsupial structure in the inner
gills. Glochidia (See Ortmann (19116, PL 89, fig. 4) measurements: 0.15 X 0.15
mm., and Surber (1915, p^ 7, fig. 9) measurements: 0.16 X 0.16 mm.).

Breeding season: Gravid females have been found on May 13, 1911; May 23,
1909; May 23, 1911; May 23, 1912; May 24, 1911; June 18, 1908; June 22,
1908; June 23, 1910; July 4, 1909; July 8, 1909; July 19, 1909. Glochidia have
been found on June 18, June 23, and July 4. Thus this form is tachytictic, breed-
ing from the middle of May to the middle of July, and beginning to discharge the
glochidia in the second half of June.

Remarks: This form has been hitherto regarded as a good species. In its
typical environment (small streams) it is, indeed, quite constant, although it
varies a good deal. The shape may be different, from subrotund to subovate, or
sub triangular, and quite often the lower posterior end is drawn out and even
deflected, thus producing an outline recalling that of P. ohliquum rubrum. Gen-
erally, there is no trace of a radial furrow, and the lateral faces are rather flat,
only slightly convex, and without a posterior ridge. In obesity, this form varies
greatly, and more swollen shells (See No. 2, under measurements) gradually pass
into P. ohliquum catillus, and this takes place in the downstream direction. In the
upper Allegheny occasional individuals turn up, having the dimensions of catillus;
such become more abundant in Armstrong Co., where they outnumber the coc-
cmeriw-forms. Here also individuals turn up, which show the beginning of a
radial furrow, which then lead to specimens with well-developed furrow, represent-
ing typical ohliquum.

There is complete intergradation between these forms, and it is impossible
to separate them except by drawing an artificial line.

Below Pittsburgh, in the Ohio, only a single coccineu7n has been found. Con-
ditions similar to those in the Allegheny prevail in the Beaver-drainage.

The nacre of coccineum is of a reddish tint more frequently than is the case
in any of the preceding forms.

Localities in Pennsylvania, represented in the Carnegie Museum:

Ohio proper and Beaver-drainage :

Ohio River, Cooks Ferry, Beaver Co.

80

MEMOIRS OF THE CARNEGIE MUSEUM.

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Ellwood City, Lawrence Co. (G. H. Clapp & H. H. Smith; G. L. Simpson, Jr.);
Harmony, Butler Co.

Slipperyrock Creek, AVurtemberg, Lawrence Co.

Neshannock Creek, Eastbrook and Volant, Lawrence Co.; Leesburg, Mercer Co.

Mahoning River, Mahoningtown (Type locality), and Coverts, Lawrence Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsville, Clarksville, Shenango, and
Jamestown, Mercer Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

A llegheny-dramage :

Allegheny River, Aladdin, Godfrey, Kelly, and Templeton, Armstrong Co.; Tionesta and Hickory,
Forest Co.; Warren, AVarren Co.; Larabee, McKean Co.

Buffalo Creek, Harbison, Butler Co.

Conemaugh River, New Florence, AA^estmoreland Co.

Loyalhanna River, Idlepark and Ligonier, AA'estmoreland Co.

Crooked Creek, Rosston, Armstrong Co.

Little Mahoning Creek, Goodville, Indiana Co.

Sandy Creek, Sandylake, Mercer Co.

French Creek, Utica, A^enango Co.; Cambridge Springs, Crawford Co.

Conneaut Outlet, Conneautlake, Crawford Co.

Conneauttee Creek, Edinboro, Erie Co.

Leboeuf Creek, AA^aterford, Erie Co.

Brokenstraw Creek, Garland, AA^arren Co.

Connewango Creek, Russell, AA'arren Co.

M onongahela-drainage :

Dunkard Creek, Mount Morris, Greene Co.

Lake Erie-drainage :

Conneaut Creek, Springboro, Crawford Co.; AA^est Springfield, Erie Co.®'*

Other localities represented in the Carnegie Museum:

Lake-drainage:

Shiawassee River, Genesee Co., Michigan (B. AA^alker, donor) (Saginaw-drainage).

Raisin River, Tecumseh, Lenawee Co., and Grape P. 0., Monroe Co., Flichigan (C. Goodrich).

St. Marys River, Rockford, Mercer Co., Ohio (C. Goodrich).

Swan Creek, Toledo, Lucas Co., Ohio (C. Goodrich).

Ohio-drainage:

Mahoning River, Leavittsburgh, Trumbull Co., Ohio (Smith collection).

Tuscarawas River, Ohio (Holland collection).

Scioto River, Ohio (Hartman collection).

AA" abash River, Geneva, Adams Co., and Bluffton, AA^ells Co., Ind. (C. Goodrich).

AA'est Fork River, Lynch IMlnes, Harrison Co., AA^est Aurginia; AA'est Milford, Harrison Co., AA^est AArginia
(AAh F. Graham); Lightburn and AA^eston, Lewis Co., AAust Aurginia.

The specimens from Conneaut Creek, which flows to Lake Erie, are all typical, medium-sized
coccineum, with red nacre, which color is never found in the lake-form, and are not the latter {pauper-
culum).

OKTMANN: monograph of the naiades of PENNSYLVANIA.

81

Little Kanawha River, Grantsville, Calhoun Co. (W. F. Graham) ; Burnsville, Braxton Co., West Virginia.
Little Coal River, Boone Co., West Virginia (Hartman collection).

Licking River, Farmer, Rowan Co., Kentucky.

Tennessee-drainage :

Holston River, Hodges, Jefferson Co.; Noeton, Grainger Co., Tennessee.

Clinch River, Solway, Knox Co., Tennessee.

West of Mississippi:

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

White River, Garfield, Benton Co., Arkansas (R. L. Moodie).

Distribution and Ecology in Pennsylvania (See fig. 8) : In Pennsylvania, this
form is widely distributed. It is especially abundant in the Beaver-drainage,
and also in that of the upper Allegheny, going up far into the headwaters, reaching
into Beaver and French Creek, Crawford and Erie Cos., and in the uppermost Alle-
gheny, AIcKean Co. In the Monongahela-drainage, it is rare, at least in Penn-
sylvania, and found only in Dunkard Creek, but formerly it surely existed else-
where in smaller creeks, which are now polluted. It becomes more abundant
again in the headwaters of the Alonongahela in West Virginia.

It is distinctly a form of the smaller creeks, and locally quite plentiful in them.
It prefers sand and fine gravel, in or below riffles, where the current is rather
lively, and generally it is deeply buried. It avoids coarse gravel, but may be found
now and then in rough places.

As has been stated, farther down in the larger rivers, it passes into P. ohliquum
catillus, and with the change of the shape of the shell, there is also a change in its
ecological preferences, or perhaps it might be true to say that the latter very likely
is the cause of the former change. We have here a good example of the correlation
of habitat and shape. P. ohliquum coccineum is distinctly an ecological race of
P. ohliquum (and its variety catillus) characteristic of the fine gravel and sand
of the smaller streams. In the Ohio below Pittsburgh coccineum is practically
absent, only a single individual having been found.

Its presence in Conneaut Creek, a tributary of Lake Erie, should be noted.
It appears that this form is not derived from the form found in Lake Erie {yau-
yerculum), but is identical with the form present in the upper Beaver and French
Creek-drainages. A crossing over the divide is in this case suggested, and it should
not be forgotten, that the old Beaver Canal (now obliterated) once connected
these systems.

General distribution: Type locality, Mahoning River, near Pittsburgh (Conrad)
(See above, p. 78, footnote 53).

Simpson (1900) gives for this form: “entire upper Mississippi-drainage,”

82

MEMOIRS OF THE CARNEGIE MUSEUM,

while Call (1885) says: “from western New York to Kansas; south to the Holston
River, Tennessee.”

Very likely, this form is present in western New York in the Allegheny-drain-
age, judging from its presence in Pennsylvania both in Warren and McKean Cos.,
although Marshall (1895) reports it only from I^ake Erie, no doubt referring to
the variety pauperculum. In Ohio it is found all over the state (Sterki, 1907a),
but definite reports are mainlj^ from smaller streams. It is known from the
Mahoning River (Lea, Dean, 1890), Tuscarawas River (Dean, Sterki),, Scioto
River, and from Columbus (Lea). As in Pennsylvania so in Ohio it crosses
over into the lake-drainage, and is found in the Cuyahoga (Dean) and Mau-
mee-drainage (Carnegie Museum). It occurs also in the lake-drainage in
southern Michigan (Walker, 1898). It is common in Indiana, and there also
crosses into the lake-drainage, St. Marys and St. Josephs Rivers (Call, 1900).
Y'ilson & Clark (1912, p. 42) observe that coccineum is found in the headwaters of
the Kankakee River in Indiana, but that in the lower Kankakee and in the Iroquois
River in Illinois it passes into a form belonging to the o5h’gaam-group. In Illinois
the same conditions seem to prevail (Baker, 1906) or at any rate it goes far up
toward the north in that state.

As our localities show, this form is found in the tributaries of the Ohio in
West Virginia and eastern Kentucky, Other records from Kentuck}^ and from
Tennessee are missing, except Call’s statement, that it is found in the Holston.
It is, indeed, present in the Holston and in the Clinch, but is quite rare there, and
closely connected with the variety catillus.

Farther to the west and southwest there is the record from Coon River, Dallas
Co., Iowa (Alarshall, 1895), that from the drainage of Turkey, Wapsipinicon, and
Volga Rivers, northeastern Iowa (Geiser, 1910), and that from Missouri (Utter-
back, 1916). It is said to be present in Kansas (Scammon, 1906), and is in northern
Arkansas (see our specimens and Meek & Clark, 1912) and even in the Ouachita
River, in Clark Co. (Wheeler, 1918). But in this direction, the limit of its range
is obscure. It seems, that in this region likewise it prefers smaller streams, but
this needs further investigation.

Lea gives it from the Ohio at Marietta, Washington Co., Ohio, and it is in the Cincinnati-list
(Harper, 1896); but these records are to be doubted. I have never found it in the Ohio, and the large
shell-heaps of the clam-diggers examined by myself at various places never yielded a single individual of
this shell,

ORTMANN: monograph of the naiades of PENNSYLVANIA.

83

Pleurobema obliquum pauperculum (Simpson) (1900).

Quadrula coccinea paupercula Simpson, 1900, p. 789; Quadrula subrotunda Sterki,
1907a, p. 391 (partim); Quadrula subrotunda Ortmann, 19096, p. 203 {per
errorem)] Quadrula coccinea magnalacustris Simpson, 1914, p. 884.®®

Plate VII, fig. 6.

Records from Pennsylvania :

Ortmann, 19096, p. 203 (as form of Q. subrotunda).

Characters of variety: Resembling P. obliquum coccineum, but shell consider-
ably smaller, more convex (diameter ranges around fifty percent), with the outline
subovate, and generally more elongate, and the growth-lines more distinct and
quite regular. The beaks are not very prominent (hence the subovate, and not
triangular, outline), and a furrow, except in extreme cases is absent, there being
merely a flattening in its place. Nacre always white.

L.

H.

D.

Pr.ct.

1.

Erie, Cat. No. 61.4393 (largest at hand). . .

. .71 mm.

50 mm.

35 mm.

.49

2.

Erie, Cat. No. 61.4393

..70 “

55 “

34 “

.49

3.

Erie, Cat. No. 61.4579

..62 “

48 “

28 “

.45

4.

Erie, Cat. No. 61.3930 (smallest at hand) .

..55 “

47 “

29 “

.53

Soft parts: Only two individuals have ever been found alive by myself, of
which the soft parts of one were preserved. It proved to be a male, with the
structure agreeing with that of P. obliquum. The characteristic features of the
females have not been observed. The color of the soft parts was grayish-white.

Glochidia and breeding season unknown.

Remarks: As Simpson correctly recognized, this is a form derived from coc-
cineum, and not directly from obliquum: nevertheless, the rules of nomenclature
force us to use the above name.

This is the Lake Erie form of coccineum, descended from the stock which
crossed the divide between the upper Ohio and the lake-drainages in northern
Indiana, Ohio, and possibly Pennsylvania. This lacustrine form has the distinct
and regular growth-lines, which are also more closely set, found so often in shells
from Lake Erie, and this peculiarity together with its small size are its chief diag-
nostic characters. In shape it is generally more elongate-ovate than coccineum,
but, according to the growth-lines, young specimens must be shorter and more
rounded-ovate. On the average it is slightly more swollen than coccineum, but

The change of the varietal name introduced by Simpson in 1914 is unnecessary. There is a
Unio pauperculus Lea (1861) but this does not constitute a pre-occupation of the term Quadrula coccinea
paupercula.

84

MEMOIRS OF THE CARNEGIE MUSEUM.

the difference in this respect is not great. There is much variation in shape,
chiefly witli regard to the elongation of the shell and the position of the beaks.
In some specimens the beaks are much anterior, thus imitating the condition seen
in P. ohliquum rubruni.

The color of the epidermis is generally dark brown, but specimens from Michi-
gan arc light brown, with greenish capillary rays; however, these specimens are
greatly water-worn.

I have not seen any specimens grading toward the parent-form, and for this
reason, we might perhaps be justified in regarding this as a true species. Never-
theless I hesitate to take this course, since I consider my material (about fifteen
specimens) as insufficient to settle this question.

Localities represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania.

Lake Erie, La Plaisance Bay, Monroe Co., iMichigan (C. Goodrich).

Distribution and Ecology (Sec fig. 8): Type locality, St. Lawrence basin at
Niagara Falls (Simpson).

Sterki mentions this form, as a dwarf race of Q. subrotunda, from the Ohio
shores of Lake Erie. Walker gives it (1913, p. 29) from Lake Erie, and probably
his Q. subrotunda refers to the same form. I have found it in Presque Isle Bay,
and received specimens from La Plaisance Bay. Probably, Marshall’s (1895)
U. coccineus from Erie Co., New York, is also this.

Our present knowledge thus limits its range to Lake Erie and the Niagara
River just below it. Simpson, indeed, says '‘and tributaries,” but exact localities
are not given.

In Presque Isle Bay I found this shell only on the North shore in a few feet
of water upon pure sand, but most of my specimens were dead shells. It is decidedly
rare at this locality.

Pleurobema obliquum rubrum (Raflnesque) (1820).

Quadrula pyramidata (Lea) Simpson, 1914, p. 888; Quadrula rubra (Rafinesque)
Vanatta, 1915, p. 557.

Plate VI, fig. 7.

Records from Pennsylvania :

Harn, 1891 '(as mytiloides) (western Pennsylvania).

Stupakoff, 1894 (Allegheny Co.).

Ortinann, 19096, p. 199 (under ohliqua).

Characters of variety: Like P, obliquum, but shell very oblique, with the beaks

ORTMANN: monograph of the naiades of PENNSYLVANIA.

85

strongly procurved, and situated at the anterior end of the shell. Radial furrow
more or less distinct. Diameter generally considerably over fifty jiercent of the
length, but sometimes less. Nacre typically red, but sometimes pinkish or white.

L.

H.

D.

Pr.ct.

Size: 1.

Industry, Cat. No. 61.3896

91 mm.

73 mm.

54

mm. .59

2.

Kelly, Cat. No. 61.3079

86 “

71 “

47

“ .55

3.

Godfrey, Cat. No. 61.3891

63 "

53 “

36

“ .57

Soft parts (See Ortmann, 1912, p. 264) : Glochidia not yet observed.

Breeding season: Gravid specimens are at hand collected on May 25, 1914 and
July 28, 1913. They all had eggs only.

Remarks: There is not the slightest doubt that this is in Pennsylvania only
an individual variation of P. obliquum, characterized by excessive obliquity.
There is great variation in this respect, and complete connection with the main
form. While the nacre is mostly red in typical specimens from the Tennessee,
in Pemisylvania it is more frequently pink or whitish. Altogether, typical ruhrum
is quite scarce in our state, but, as our fig. 7 on PL VI shows, it is present, and does
not differ at all from specimens from the Tennessee. The intergrades toward
P. obliquum are much more abundant with us.

Considering the conditions in Pennsylvania alone, I should never have thought
of giving this form varietal rank. _ But in other regions, it is more distinctly marked,
as for instance in the Upper Tennessee-drainage. Here the var. rubrum in its
typical phase is quite abundant and rather sharply distinguished. However,
here also are found forms representing transitions to obliquum and catillus. In
the region west of the Mississippi, where P. obliquum is practically missing, the
var. rubrum (and catillus) are found in its place, and thus it seems to be advisable
to retain rubrum as a distinct variety. These western forms require still closer
study.

I want to emphasize this instance, for it demonstrates that a certain form may
be in a certain region an individual variation of another form, while in other regions
it becomes a better defined local race, or variety.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Shipping-port and Industry, Beaver Co.; Neville Island, Allegheny Co.

Allegheny River, Godfrey and Kelly, Armstrong Co.

Other localities represented in the Carnegie Museum:

Ohio-drainage:

Ohio River, St. Marys, Pleasants Co., West Virginia; Portsmouth, Scioto Co., Ohio.

Tuscara-was River, Ohio (Holland collection).

West Fork of White River, Riverside, Greene Co., Indiana (J. D. Haseman).

86

MEMOIRS OP THE CARNEGIE MUSEUM.

Tennessee-drainage :

Tennessee River, Knoxville, Knox Co., Tennessee.

French Broad River, Boyd Creek, Sevier Co., Tennessee.

Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill, Noeton, and
Holston Station, Grainger Co.; Austin Mill, Hawkins Co., Tennessee.

Clinch River, Solway, Knox Co.; Edgemoor and Offutt, Anderson Co.; Needhams Ford (B. Walker,
donor) and Black Fox Ford, Union Co.; Clinch River Station, Claiborne Co.; Oakman, Grainger
Co., Tennessee.

There are a good many specimens at hand from west of the Mississippi, from Arkansas, Kansas,
and Oklahoma, but since these form a peculiar and rather obscure association of forms, partly intergrading
with the var. catillus, partly having characters of their own, I omit them here, till further investigations
have been made.

Distribution in Pennsylvania (See fig. 8) :

The range of this form in Pennsylvania, is coextensive with that of P. ohliquum,
but it is rather rare. The same is true farther down the Ohio, and wherever I
collected it myself between Pittsburgh and Cincinnati I found only a few specimens
of it among large numbers of ohliquum and its intergrades.

General distribution: Type locality, Kentucky River (Rafinesque) .

Where P. ohliquum is found, this form generally is also present. However,
P. ohliquum ruhrum seems to have a wider range than the main form, for it is also
found west of the Mississippi without being associated with P. ohliquum (See
above). I also have observed in the upper Tennessee-drainage, that it ascends
in the Holston and Clinch a little farther than the main species.

Note: Attention should be called again to the fact, that we have here a group
of forms (obliquum-group) , of which some have swollen shells {ohliquum and catillus),
which are found in the larger rivers, while another allied form has a compressed
shell {coccineum), which is peculiar to small creeks. As in certain other cases,
there is also a dwarfed lacustrine form, pauper culum.

It is well to keep these facts in mind. Aly observations on the passing of the
creek form into that of the large rivers have been confirmed for the Kankakee-
drainage in Indiana and Illinois by Wilson & Clark (1912, p. 42). However, the
remark of these authors, that I am of the opinion that these forms are “identical’’
rests upon a misunderstanding. They are “conspecific,” and represent well
marked races, or varieties, of the same species, but they are not identical.

Pleurobema clava (Lamarck) (1819).

Pleurohema clava (Lamarck) Simpson, 1914, p. 735.

Plate VH, figs. 7, 8, 9.

Records from Pennsylvania :

Call, 1885 (Allegheny River, as U. patulus).

Harn, 1891 (western Pennsylvania).

OETMANN: monograph of the naiades of PENNSYLVANIA.

87

Marshall, 1895 (Allegheny River, Warren Co.).

Ortmann, 19096, p. 197.

Characters of the shell: Shell small, or barely of medium size, but comparatively
heavy. Outline subtriangular to subovate, more or less elongated and drawn out
posteriorly, very oblique, often cuneiform (high in front, tapering behind) . Lower
margin convex or straight. Beaks not much elevated, moderately swollen, directed
obliquely forwards, and situated near the anterior end of the shell, sometimes
quite anterior. Beak-sculpture rudimentary, indistinct, consisting of three to
four subconcentric ridges. Shell more or less swollen anteriorly, less swollen and
more compressed posteriorly ; sometimes with the anterior swelling marked off quite
sharply from the posterior fiat part, but hardly ever with a distinct radial depression.
In most cases the swollen anterior part passes insensibly into the compressed
posterior part. In some cases the shell is very little swollen anteriorly, and the
valves are rather regularly and gently convex. No distinct posterior ridge. Sur-
face of shell smooth, without sculpture.

Epidermis yellowish to light brown or pale chestnut, darker when old; gen-
erally with more or less distinctly green capillary rays, which are interrupted and‘
much broken, forming very often, and characteristically, squarish, or even trans-
verse green to blackish spots or blotches. Growth-rests not very different in color.

Hinge well-developed. Pseudocardinals divergent, moderately large, ragged.
Interdentum not very wide, often rather narrow. Lateral teeth rather long.
Beak-cavity not deep. Dorsal muscle-scars on the hinge-plate. Nacre always

white.

No sexual differences in the shell.

L. H. D.

Size: 1. Harbor Bridge, Cat. No. 61.3815 (largest) 78 mm. 51 mm. 36 mm.

2. Eastbrook, Cat. No. 61.3335 -..74 “ 48 “ 27 “

3. Eastbrook, Cat. No. 61.3335 64 “ 40 “ 23 “

4. Eastbrook, Cat. No. 61.3335 54 “ 35 “ 21 “

5. Mosgrove, Cat. No. 61.4328 39 “ 29 “ 19 “

Soft parts (See Ortmann, 1912, p. 264, fig. 9). Glochidia (See Ortmann, 19116,
PL 89, fig. 5).

Breeding season: The following records are at hand for gravid females: May
22, 1912; May 23, 1911; May 23, 1912; May 24, 1911; June 18, 1908; June 27,
1910; July 8, 1909; July 8, 1911; July 10, 1908; July 19, 1909. Glochidia have
been found as early as June 18. Tachytictic form, breeding from about the middle
of May to the end of July.

Remarks: A very distinct species, at least in Pennsylvania. It belongs to

f

88

MEMOIRS OF THE CARNEGIE MUSEUM.

the smaller mussels, and its chief characters are the peculiar, obliquely-ovate,
elongated or cuneiform outline, and the light color of the epidermis, with green
patches, which are frequently (but not always) present. In shape, situation of
beaks, and obesity, it varies greatly (compare, for instance the figure of Lea’s
patulus, Obs. I, 1834, PL 12, fig. 20, and of Conrad’s clava, Mon. 1835, PL 5, fig. 1).
In old specimens the beaks are sometimes quite anterior, forming the foremost
point of the shell, but such extremes are rare in Pennsylvania, and generally the
beaks remain a little behind the anterior margin of the shell. Also with regard to
the convexity of the valves no extreme cases have been observed in Pennsylvania;
on the contrary the tendency is more toward the flatter types (corresponding more
nearly to patulus).

The peculiar color-markings are present in most specimens, but extremely
variable. The spots are found chiefly towards the beaks, and often an irregular

• Pleurohema clava.

series of these spots runs down just in front of the posterior slope. In old shells,
the epidermis tends to become uniformly brown.

Localities in Pennsylvania, represented in the Carnegie Museum:

Ohio-drainage :

Little Beaver Creek, Cannelton, Beaver Co. (Miss Vera White).

Raccoon Creek, Raccoon Township, Beaver Co.

Beaver-drainage :

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Harmony, Butler Co.

Mahoning River, Mahoningtowm and Coverts, LawTence Co.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

89

Neshannock Creek, Eastbrook, Lawrence Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Clarksville, Shenango, and Jamestown,
Mercer Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

A llegheny-drainage :

Allegheny River, Aladdin, Godfrey and Mosgrove, Armstrong Co.; Walnut Bend, Venango Co.; Tio-
nesta and Hickory, Forest Co.

Buffalo Creek, Harbison, Butler Co.

Conemaugh River, New Florence, Westmoreland Co.

Loyalhanna River, Idlepark and Ligonier, Westmoreland Co.

Sandy Creek, Sandylake, Mercer Co.

French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co.
Conneaut Outlet, Conneautlake, Crawford Co.

Leboeuf Creek, Waterford, Erie Co.

Monongahela-drainage:

Dunkard Creek, Mount Morris, Greene Co.

Cheat River, Cheat Haven, Fayette Co.

Other localities represented in the Carnegie Museuyn:

Lake-drainage:

Maumee River, Defiance, Defiance Co., Ohio, and Fort Wayne, Allen Co., Indiana (C. Goodrich).
Ohio-drainage:

Tuscarawas River, Ohio (Holland collection).

Wabash River, Geneva, Adams Co., Indiana (C. Goodrich).

West Fork River, Lynch Mines, Harrison Co.; Lightburn and Weston, Lewis Co., West Virginia.

Little Kanawha River, Burnsville, Braxton Co., West Virginia.

North Fork Hughes River, Cornwallis, Ritchie Co., West Virginia.

Elk River, Sutton and Gassaway, Braxton Co.; Shelton, Clay Co., West Virginia.

Tennessee-drainage :

Tennessee River, Tuscumbia, Colbert Co., Alabama (H. H. Smith).

Distribution and Ecology in Pennsylvania (See fig. 9) : The range of this species
in Pennsylvania is very similar to that of P. obliquum coccineum, with the exception
that it does not go into the lake-drainage. The two forms are very often found
associated. P. clava does not go up the Allegheny as far as coccineum does.”

Altogether, P. clava is a rare shell, and never found in great numbers. It is
found mostly in sand and fine gravel, and is deeply buried. In the large rivers it is
missing. The lowest point in the Allegheny is at Aladdin. It has never been
found in the Monongahela proper and in the Ohio, and is also missing in the lists
of the older collectors (Stupakoff, Clapp, Rhoads). .The same holds good farther
down the Ohio. I have never seen a trace of it between Pittsburgh and Cincinnati.
General distribution: Type locality, “Lake Erie” (Lamarck).

” I found P. clava as far up as Hickory, Forest Co., but Marshall (1895) reports it from Warren Co.

90

MEMOIRS OF THE CARNEGIE MUSEUM.

We have here the curious fact that a species is not found at its type locality.
It should lie noted, however, that this species is present in tributaries of the lake,
chiefly in the Maumee (Goodrich, 1914) substantiated by specimens in the Carnegie
Museum.

Although Call (1895 and 1900) cites western New York and the Ottawa
River, Canada, for this species, there is serious doubt whether it goes so far north
and northeast. No exact localities are known in the Allegheny River in the state
of New York, nevertheless this species might be there, since it has been reported
from Warren Co., Pennsylvania, and is positively found immediately below in
Forest Co. Its main range includes the Ohio, Cumberland, and Tennessee systems,
chiefly in their tributaries. It is widely distributed in these drainages in western
Pennsylvania, Ohio, Indiana, West Virginia, Kentucky, Tennessee, and northern
Alabama. In Ohio and Indiana it surely crosses over into the lake-drainage, St.
Marys and Maumee Rivers (Call, 1900, and Sterki, 1907a, also collected by C.
Goodrich). Nevertheless it has not been reported from Lakes Erie and Michigan
(See Walker, 1898 and 1913). Remarkably enough, while it is widely distributed
in Indiana (Call, 1896a and 1900), it is listed from Illinois (by Baker, 1906) only
from the Wabash. Its actual absence in the rest of Illinois is confirmed by the
recent investigations of Wilson and Clark. According to them it is in Tippecanoe
Lake (Wabash-drainage) in Indiana, but not in the Kankakee-drainage in Indiana
and Illinois. West of Illinois, only three records are at hand (from Iowa City,
Iowa; St. Peters River, Minnesota; and Nebraska), which, however, are very
likely incorrect (Simpson, 1900, p. 746 seems to have no confidence in them, and
Geiser (1910) does not give this shell from northeastern Iowa). No other records
are known from west of the Mississippi.

In the Tennessee-drainage, this species surely goes to northern Alabama.
In the upper Tennessee region it is missing but there are a number of closely allied
species or forms, the standing of which will be elucidated elsewhere; also in the
Coosa-Alabama-drainage there are representatives, which have been taken for
synonyms, for instance by Call. But this surely is incorrect, and requires further
study.

It is not known, whether this species outside of Pennsylvania inhabits pre-
ferably the smaller streams, except that it is surely present in them in West Virginia.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

91

Genus Elliptio Rafinesque (1820).

Ortmann, 1912, p. 265; Simpson, 1914, p. 586 (as section of Unio).

Type Unio nigra Rafinesque.

Five species and one variety are found in Pennsylvania (one of them somewhat
doubtful as to its presence in the state).

Key to the Pennsylvanian Species and Variety of Elliptio.

Hi. Shell very large, rhomboid-ovate, not elongated, with a sharp posterior ridge. Posterior slope (at

least in the young) with radiating wrinkles. (Western shell.) E. niger.

a 2- Shell larger or smaller, elongated ovate, or subtrapezoidal, with ill defined or no posterior ridge.
Posterior slope not wrinkled.

bi. Shell rather large, elongated ovate, heavy, not subtrapezoidal, somewhat inflated or subcom-
pressed. (Western shells.)

Cl. Shell larger, subcompressed. Epidermis dark colored, with obscure growth-lines.

E. dilatatus-

C2. Shell smaller, more swollen and thus more nearly subcylindrical. Epidermis of lighter

color, with more distinct and more regular growth-lines E. dilatatus sterkii.

6 2. Shell moderately large or small, subtrapezoidal, more or less elongated, and more or less com-
pressed. (Eastern shells.)

Cl. Shell moderately large, subtrapezoidal, slightly elongated, not much pointed behind.

E. violaceus.

C2. Shell moderately large or small, subtrapezoidal, but much elongated, so as to become lance-
shaped, pointed behind.

di. Shell rather small, upper and lower margins nearly jiarallel; greatest height in the

middle portion. . , E. cupreus.

d2- Shell somewhat larger, upper and lower margins not subiiarallel ; greatest height more
anteriorly E. fisherianus

Elliptic niger (Rafinesque) (1820).

Unio crassidens Lamarck. Simpson, 1914, p. 606.

Plate VIII, fig. 1.

Records from Pennsylvania :

Call, 1885 (Allegheny River).

Harn, 1891 (western Pennsylvania).

Clapp, 1895 (Allegheny Co.).

Marshall, 1895 (Allegheny River, Warren Co.).®®

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.).

Ortmann, 19095, p. 197.

Characters of the shell: Shell large and very heavy. Outline ovate-rhomboid,
rather variable, shorter or longer, but always longer than high, but rarely almost

®® This record is doubtful. I have never seen this striking shell in the Allegheny above Oil City.

92

MEMOIRS OF THE CARNEGIE MUSEUM.

twice as long as high. Anterior end rounded, lower posterior end more or less
produced and bluntly pointed. Lower margin convex, straight, or even sometimes
slightly concave in its posterior part. Beaks not greatly swollen and not very
prominent, situated in advance of the middle of the shell, but not very near the
anterior end. Beak-sculpture very rudimentary, almost obliterated, consisting of
a few weak concentric ridges running parallel with the growth-lines. Shell moder-
ately swollen towards the beaks, but sides of the disk only gently convex, often
quite flat posteriorly. A distinct and rather sharp posterior ridge runs from the
beaks toward the lower posterior end. The posterior slope behind this ridge is
subtruncate and fiat, and toward the beaks it is generally ornamented with fine
radiating wrinkles or corrugations. The latter are chiefly seen in young shells,
are irregular and often interrupted. Upon old shells they are absent, mostly
destroyed by the erosion of the beaks; sometimes they occur also in young speci-
mens poorly developed, or are absent. Otherwise the surface of the shell is smooth,
without sculpture.

Epidermis brown, from reddish brown to dark blackish brown; in the young
it is lighter brown, and generally marked by dark green, obscure rays, which are
narrower or wider, but in adult shells the rays disappear entirely. Growth-rests
not distinctly marked by color.

Hinge-teeth well-developed and very strong. Pseudocardinals divergent,
large, and ragged. Interdentum rather narrow. Lateral teeth thick and rather
long. Beak-cavity not deep. Dorsal muscle-scars on the hinge-plate. Nacre
always some shade of red. Very rarely it is almost white in the centre of the shell,
and then some color is always present around the margins. In most cases, the
whole nacre is tinted, and the color ranges from a beautiful salmon through all
shades of pink and red to a blueish purple.

No sexual differences in the shell.

L. H. D.

1. Kelly, Cat. No. 61.3777

129 mm.

85

mm.

49

mm.

2. Cooks Ferry, Cat. No. 61.4427

125 “

87

ii

57

3. Kelly, Cat. No. 61.3055

117 “

73

a

44

H

4. Neville Island, Cat. No. 61.926

119 “

85

ii

56

a

5. Kelly, Cat. No. 61.3059

89 “

60

43

u

6. Shippingport, Cat. No. 61.4607

63 “

42

a

23

Soft parts (See Ortmann, 1912, p. 266, fig. 10). Glochidia (See Ortmann,
19116, PL 89, fig. 6) not quite mature, size: 0.13 X 0.15 mm. Mature glochidia
have been described and figured by Surber (1915, p. 8, fig. 13). Their shape is
somewhat pointed (subtriangular) ; their size: 0.15 X 0.16 mm.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

93

Breeding season: Gravid females have been found only once by myself, on
June 22, 1909 (with eggs and young glochidia). Surber secured them on July 14,
1911. Thus the duration of the breeding season remains uncertain, but probably
is short, and ends in July.

Remarks: A very characteristic species, which cannot be confused with any
other form. In fact Rafinesque’s definition of U. nigra as a ^Targe, heavy shell.

Fig. 10.

■ Elliptio niger.

□ do. do. (Indian garbage heap.)

• Elliptio dilatatus.

+ Elliptio dilatatus sterkii.

with red nacre, from the Ohio,” is entirely satisfactory. The shell varies greatly
in shape, and may be shorter or longer in proportion to height, but is always recog-
nized by its subovate outline, the distinct posterior ridge, and the color of the
nacre. The peculiar sculpture of the posterior slope is well-developed only in
young shells.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Smiths Ferry (W. F. Graham), Shippingport, Cooks Ferry, Industry, and Beaver, Beaver
Co.; Shoustown (R. Foerster), Coraopolis (S. N. Rhoads), and Neville Island, Allegheny Co.
Allegheny River, Harmarville and Natrona, Allegheny Co.; Braeburn, Westmoreland Co.; Aladdin,
Godfrey, Johnetta, Kelly, and Templeton, Armstrong Co.

Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).

94

MEMOIRS OF THE CARNEGIE MUSEUM.

Other localities represented in the Carnegie Museum:

Ohio-drainage:

Ohio River, Congo, Hancock Co., West Virginia; Toronto, Jefferson Co., Ohio; St. Marys, Pleasants
Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland, Meigs Co., Ohio; Ports-
mouth, Scioto Co., Ohio.

West Fork White River, Riverside, Greene Co., Indiana (J. D. Haseinan).

Elk River, Gassaway, Braxton Co., and Shelton, Clay Co., West Virginia.

Levisa Fork, Big Sandy River, Prestonsburg, Floyd Co., Kentucky.

Tennessee-drainage :

Tennessee River, Florence, Lauderdale Co., Alabama (II. 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.; Noeton, Grainger Co.; Austin
hlill, Hawkins Co., Tennessee.

South Fork Holston River, Pactolus, Sullivan Co., Tennessee.

Clinch River, Solway, Knox Co.; Edgemoor, Clinton, and Offutt, Anderson Co.; Clinch River Station,
Claiborne Co.; Oakman, Grainger Co., Tennessee; Clinchport, Scott Co., Virginia.

Emory River, Harriman Junction, Roane Co., Tennessee.

Gidf-drainage:

Pearl River, Mississippi (Juny collection).

Black Warrior River, Squaw Shoals, Jefferson Co., Alabama (H. H. Smith).

Cahaba River, Pratts Ferry, Bibb Co., and Gurnee, Shelby Co., Alabama (H. H. Smith).

Coosa River, Wetumpka, Elmore Co.; Weduska and Peckerwood Shoals, and Wilsonville, Shelby Co.;

Coosa Valley and Riverside, St. Clair Co., Alabama (H. H. Smith).

Choccolocco Creek, Jackson Shoals, Talladega Co., Alabama (H. H. Smith).

Chattooga River, Cedar Bluff, Cherokee Co., Alabama (H. H. Smith).

Etowah River (R. E. Call) and Oostanaula River (G. H. Clapp, donor), Rome, Floyd Co., Georgia.
Sepulga River, Herbert, Conecuh Co., Alabama (Alabama Museum, donor).

Distribution and Ecology in Pennsylvania (See fig. 10) : This is a species re-
stricted to the large rivers, Ohio, Allegheny, and Monongahela, but extremely
abundant in them. I have never seen it in the’ Allegheny above Templeton in
Armstrong Co. In the Monongahela it once must have gone up above Charleroi
at least as far as the West Virginia state-line, since I have found it in an Indian
garbage-heap at Point Marion (See Ortmann, 1909c, p. 13).

Elliptio niger lives preferably in coarse gravel, often among heavy stones,
in strongly flowing water, to which habitat it is especially adapted by its heavy,
strong shell. Call (1900, p. 510) gives as its habitat “muddy bottoms,” which is

The specimens from the Coosa-drainage have in part been labeled by Walker as U. incrassatus
Lea. But I consider them to be E. niger. They reach a considerable size, while the real U. incrassatus
from the Chattahoochee River is a dwarf form.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

95

not at all true for Pennsylvania. In the Ohio this shell is also found in great
numbers on the shell-banks, consisting of masses of dead and broken shells, the
interspaces filled with sand and gravel, in strong, steady currents. The same is
the case farther down the Ohio, where it is taken in immense numbers by the
clam-diggers, but rejected on account of the color of its nacre.

General distribution: Type locality, Ohio River (Rafinesque).

A common shell all along the Ohio from western Pennsylvania through Ohio,
West Virginia, Kentucky, Indiana, and Illinois. Also in the Mississippi in Illinois,
Iowa, and North to Minnesota, Winona Co. (See Holzinger, 1888). It goes only
into the larger tributaries. In Ohio, it is in the Muskingum River at Marietta,
Washington Co. (Hildreth, 1828), but not in the Tuscarawas River; it occurs in
the Scioto River (Sterki, 1907a). In Indiana it is known from the Wabash (Call,
1900, Goodrich, 1914), and White River (Carnegie Museum). In Illinois it is
found in the Wabash, and also in the Kaskaskia and Spoon Rivers (Baker, 1906).
In Iowa, it is in the Wapsipinicon, at Independence, Buchanan Co. (Geiser, 1910).
In West Virginia, I have discovered it in Elk River (tributary to the Kanawha).
This river, and the Levisa Fork of the Big Sandy, are the smallest streams, in
which I have found it.

Additional records from Kentucky are missing, but this species is known from
the Cumberland, Tennessee, Duck, Holston, and Clinch Rivers in Tennessee.

In addition, it is widely distributed in the Alabama-drainage. The connection
of this range with the main range seems to be over the Gulf plain; but this requires
further study. The form from Alabama is indistinguishable from the Ohio
form, bui, judging from the material at hand, not quite so large. Eastward in
the Chattahoochee in Georgia this form passes into a still smaller form, called E.
incrassatus.

Elliptio dilatatus (Rafinesque) (1820).

Unio gibhosus Barnes. Simpson, 1914, p. 597; Unio dilatata Rafinesque,
Vanatta, 1915, p. 555; Elliptio dilatata (Rafinesque) Utterback, 1916,
p. 90.

Plate VIII, fig. 2.

Records from Pennsylvania:

Harn, 1891 (western Pennsylvania).

Stupakoff, 1894 (Allegheny Co.).

Marshall, 1895 (Allegheny River, Warren Co.).

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.; Beaver River, Wampum,
Lawrence Co.).

Ortmann, 19096, p. 197.

96

MEMOIRS OF THE CARNEGIE MUSEUM.

Characters of the shell: Shell rather large and heavy. Outline elongate-ovate,
generally about twice as long as high, and often longer. Anterior end rounded,
liosterior end produced and narrowing, but blunt. Lower margin gently convex,
straight, or even slightly concave. Upper margin curving gently down into the
upper posterior margin, without forming a distinct angle, and thus the shell is
not sub trapezoidal. Beaks not prominent, situated near the anterior end. Beak-
sculpture distinct, consisting of four to five rather heavy bars; the first two are
subconcentric, the following ones run in the direction of the growth-lines, are almost
straight in the middle, with the anterior and posterior parts obliterated; sometimes
an indication of a sinuation is seen a little behind the middle. Shell somewhat
swollen in the anterior part and toward the beaks; less swollen and almost flat,
sometimes even compressed posteriorly and toward the lower margin, without a
distinct posterior ridge. Surface smooth, without any sculpture.

Epidermis brown to black. In young specimens, traces of greenish rays are
discernible, but generally the color is rather uniform, and the growth-rests are
likewise not marked by darker color.

Hinge-teeth well-developed, but not very heavy. Pseudocardinals divergent,
ragged. Interdentum rudimentary or absent. Lateral teeth long. Beak-cavity
shallow. Dorsal muscle-scars in the beak-cavity. Nacre white or variably
colored from light pink and salmon to deep purple, and coppery purple; often
parti-colored, white and salmon, white and purple, salmon and purple, etc.

No sexual differences in the shell.

1. Jamestown, Cat. No. 61.3752 (largest

L.

H.

D.

Pr.ct.

at hand)

. . . 136 mm.

61 mm.

35 nim.

.24

2. Industry, Cat. No. 61.3763

...122 “

55 “

33 “

.27

3. Greenville, Cat. No. 61.3329

. . . 91 “

37 “

27 “

.30

4. Wampum, Cat. No. 61.2889

. . . 54 “

26 “

13 “

.24

Soft parts (See Ortmann, 1912, p. 271). The Glochidia have been figured by
Lea (Obs. XIII, 1874, PI. 21, fig. 10), by Lefevre & Curtis (1910, p. 97, fig. N, &
1912, p. 146, fig. 0) by Ortmann (19116, PI. 89, fig. 7) and Surber (1912, PL 2,
fig. 38).

Breeding season: I have found several hundreds of gravid females on numerous
dates between Alay 11 and August 13. Glochidia have been seen as early as June
5; but on the latest date (August 13) a specimen with eggs was observed. This
is clearly an abnormal case. Surber (1912, p. 7) gives the breeding season as from
June to August, but it begins before the middle of May. The species is typically
tachytictic.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

97

Remarks: I know of only two species in Pennsylvania, with which the jiresent
species might be confounded according to the general shape : Ellipsaria fasciolaris
(Rafinesque) and Eurynia recta (Lamarck). The former is easily distinguished
by the light, yellowish brown color of the epidermis, variegated with green rays and
blotches; the latter is a straighter shell, more pointed behind, with a greenish
black and shining epidermis.

Elliptio dilatatus is variable chiefly in size. In the larger rivers generally,
and sometimes also in smaller creeks, it grows to a considerable length; while in
other streams, chiefly those of the mountains, it remains rather small. Its general
shape is rather uniform: it always is an elongated shell, produced and tapering
behind, with no indication of a trapezoidal outline, and with a bluntly pointed
posterior end. Its nacre is very variable in color, but while in the Ohio and Alle-
gheny all shades are found, and shells with white nacre are quite abundant, in
other sections, as for instance in the whole Beaver-drainage, shells with light-
colored nacre are extremely rare. There is no tendency to form local races, except
that a dwarfed form prevails in the mountain-streams.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio-dramage :

Ohio River, Smiths Ferry, Shippingport, Cooks Ferry, and Industry, Beaver Co.; Coraopolis (S. N.
Rhoads) and Neville Island, Allegheny Co.

Little Beaver Creek, Cannelton (Miss Vera White), Darlington, and New Galilee, Beaver Co.; Enon
Valley, Lawrence Co.

Beaver-dramage :

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Ellwood Citj", Lawrence Co. (G. L. Simpson, Jr.).

Slipperyrock Creek, Wurtemberg and Rose Point, Lawrence Co.

Wolf Creek, Grove City, Mercer Co.

Mahoning River, Mahoningtown, Coverts, Edinburg, and Hillsville, Lawrence Co.

Neshannock Creek, Eastbrook and Volant, Lawrence Co.; Leesburg, Mercer Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsville, Clarksville, Shenango, and
Jamestown, Mercer Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

Little Shenango River, Greenville, Mercer Co.

Allegheny-drainage:

Allegheny River, Natrona, Allegheny Co.; Schenley, Aladdin, Godfrey, Johnetta, Kelly, Mosgrove,
Templeton, and Parkers Landing, Armstrong Co.; Walnut Bend, Venango Co.; Tionesta and
Hickory, Forest Co.; Warren, Warren Co.; Larabee, McKean Co. (Dennis Dally).

Loyalhanna River, Idlepark and Ligonier, Westmoreland Co.®“

Yellow Creek, Homer, Indiana Co.

“ Dead shells have been seen in the Conemaugh River, at New Florence, Westmoreland Go,

98

MEMOIRS OF THE CARNEGIE MUSEUM.

Quemahoning Creek, Stantons Mill, Somerset Co.

Crooked Creek, Rosston and South Bend, Armstrong Co.

Little Mahoning Creek, Goodville, Indiana Co,

Sandy Creek, Sandylake, Mercer Co.

French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co.
Conneaut Outlet, Conneautlake, Crawford Co.

Conneaut Lake, Cra'wford Co.

Cussewago Creek, Mosiertown, Crawford Co. (H. & L. Ellsworth).

Leboeuf Creek, Waterford, Erie Co.

Connewango Creek, Russell, Warren Co.

Potato Creek, Smethport, McKean Co. (P. E. Nordgren).

Monongahela-drainage:

Monongahela River, Westmoreland Co. (G. A. Ehrmann); Charleroi, Wasliington Co. (G. A. Ehrmann) ;

Millsboro, AVashington Co. (J. A. Shafer).

Dunkard Creek, Wiley, Dunkard, and Mount Morris, Greene Co.®‘

Cheat River, Cheat Haven, Fayette Co.

Lake Erie-drainage:

Conneaut Creek, West Springfield, Erie 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:

Grand River, Cayuga, Haldimand Co., Ontario, Canada (C. Goodrich).

Swan Creek, Toledo, Lucas Co., Ohio (C. Go'odrich).

Maumee River, Defiance, Defiance Co., Ohio (C. Goodrich).

Raisin River, Monroe and Grape P. 0., Monroe Co., Michigan (C. Goodrich).

Kalamazoo, Kalamazoo Co., Michigan (Hartman collection).

Ohio-drainage :

Chautauqua Lake, Chautauqua Co., New York.®^

Tuscarawas River, Ohio (Holland collection).

AVest Fork AA^hite River, Riverside, Greene Co., Indiana (J. D. Haseman).

AA^ abash River, Bluffton, AATlls Co., Indiana (C. Goodrich); AA^hite Co., Illinois (G. H. Clapp, donor).
Ohio River, Congo, Hancock Co., AATst Adrginia; Toronto, Jefferson Co., Ohio; St. Marys, Pleasants
Co., AATst Afirginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio.

Cheat River, Jaco, Monongalia Co., AAYst Virginia.

AATst Fork River, Lynch Alines, Harrison Co.; Lightburn and AATston, Lewis Co., AVest Virginia.

Little Kanawha River, Grantsville, Calhoun Co. (AV. F. Graham); Burnsville, Braxton. Co., AVest Virginia.
North Fork Hughes River, Harrisville (AA^. F. Graham) and Cornwallis, Ritchie Co., AA^est Virginia.

Elk River, Shelton and Clay, Clay Co.; Gassaway and Sutton, Braxton Co., AVest Virginia.

Pool of Kanawha River, Glen Ferris, Fayette Co., AVest Virginia.

Dead shells have been seen in Tenmile Creek, Amity, AVashington Co.

Particulars gbout these see below, under var, sterkii.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

99

Greenbrier River, Ronceverte, Greenbrier Co., West Virginia.

New River, Hinton, Summers Co., West Virginia; Pearisburg, Giles Co., Virginia.

Reed Creek, Wytheville, Wythe Co., Virginia.

Licking River, Farmer, Rowan Co., Kentucky.

Cumberland- and Tennessee-drainage :

Cumberland River, Orby, Bell Co., Kentucky.

Tennessee River, Tuscumbia, Colbert Co., and Florence, Lauderdale Co., Alabama (H. H. Smith).
Shoals Creek, Lauderdale Co., Alabama (H. H. Smith).

Elk River, Estill Springs, Franklin Co., Tennessee (H. H. Smith).

Flint River and Hurricane Creek, Gurley, Madison Co., Alabama (H. E. Wheeler).

Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).

South Chickamauga Creek, Ringgold, Catoosa Co., Georgia.

Tennessee River, Knoxville, Knox Co., Tennessee.

French Broad River, Boyd Creek, Sevier Co., Tennessee.

Little Pigeon River, Sevierville, Sevier Co., Tennessee.

Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill, Noeton, and
Holston Station, Grainger Co.; Austin Mill and Church Hill, Hawkins Co., Tennessee.

South Fork Holston River, Pactolus, Bluff City, and Emmett, Sullivan Co., Tennessee.

Middle Fork Holston River, Chilhowie, SmjTh Co., Virginia.

North Fork Holston River, 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 and Clinchpoi’t, Scott Co., Vir-
ginia; St. Paul, Wise Co.; Fink and Cleveland, Russell Co.; Raven, Richland, and Cedar Bluff,
Tazewell Co., Virginia.

Emory River, Harriman, Roane Co., Tennessee.

Powell River, Combs, Claiborne Co., Tennessee; Dryden, Lee Co., Virginia.

Wallen Creek, Lee Co., Virginia (G. H. Clapp, donor).

Mississippi and westwards:

Minnehaha Falls, Minneapolis, Hennepin Co., Minnesota (P. E. Nordgren).

Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

White River, Cotter, Baxter Co., Arkansas (A. A. Hinkley).

Black River (H. E. Wheeler), and Spring River (A. A. Hinkley), Black Rock, Lawrence Co., Arkansas.
Saline River, Benton, Saline Co., Arkansas (H. E. Wheeler).®^

Ouachita River, Arkadelphia, .Clark Co., Arkansas (H. E. Wheeler).®^

Neosho River, Kansas (R. L. Moodie); Miami, Ottawa Co., Oklahoma (F. B. Isely).

Wea Creek and Bull Creek, Miami Co., Kansas (C. Goodrich donor) (Osage-drainage).
Alahama-drainage:

Buttahatchee River, Hamilton, Marion Co., Alabama (H. H. Smith).

Sipsey River, Elrod, Tuscaloosa Co.; Fayette, Fayette Co., Alabama (H. H. Smith).

A peculiar race, dwarfed, nacre white, inclining toward var. subgibbosus (Lea).

Mostly the var. subgibbosus, but some are undistinguishable from the normal form. (See Wheeler,

1918.)

Alabama specimens are lai^ely the var. subgibbosus. But the specimens recorded here are typical
dilatatus (often intergrading with subgi' bosus) .

100

MEMOIRS OF THE CARNEGIE MUSEUM.

Cahaba River, Gurnee, Shelby Co., Alabama (H. H. Smith).

Distribution and Ecology in Pennsylvania (See fig. 10) : In the Ohio and Lake-
drainage of western Pennsylvania this is, next to Strophitus edentulus, the most
abundant species. It is practically ubiquitous, and there are few streams, which
contain any shells, in which it has not been found. It is moreover common in
the large rivers as well as in small creeks, and in some of the latter it is the leading
form.

This wide and universal distribution undoubtedly is due to the fact, that
this species is not very jiarticular with regard to its station, and it is hardly possible
to say that it prefers any definite ecological conditions. It is found in riffles in
small streams, as well as among heavy rocks in the larger rivers; it is upon the
shell-banks of the Ohio, as well as in quiet iiools and eddies with muddy bottom;
it is in lakes u]ion sandy, gravelly, and muddy bottoms. Nevertheless the uni-
formity of its characters is quite remarkable, and even beyond the limits of Penn-
sylvania there is hardly any change.

General distribution: Type locality, not siiecifically given by Rafinesque, but
Vanatta gives “Kentucky River.”

According to Simpson (1900), this species has the following range: “Entire
Mississippi-drainage; St. Lawrence and its tributaries; Alabama River system;
southeast into Florida; southwest to Guadeloupe River, Texas.” However, this
includes several varieties.

Typical E. dilatatus is certainly found in western New York, occurs in the
Kanawha system up to the Bluestone River in AAest Virginia, and in New River
to Virginia (See Call, 1885, and Carnegie Museum). In the Tennessee it extends
to northern Alabama, and in the headwaters to southwestern Virginia. North-
ward it is found in Canada, Michigan, Wisconsin, and Minnesota- Westward it
extends to Iowa, Kansas, Oklahoma, and northern Louisiana. In the Alabama
system and in southern Arkansas it is largely represented by the form subgibbosus
(Lea) (See Lewis, 1877). But some specimens from the Cahaba, Black Warrior,
and Tombigbee-drainages in the Carnegie Museum, represent the typical phase,
while specimens from the Coosa are all subgibbosus. Likewise in Arkansas the
two forms seem to pass into each other (See Wheeler, 1918).

E. dilatatus passes over into the drainage of Lake Michigan and Lake Erie,
but, as we shall see below, in Lake Erie proper, it assumes a peculiar form, which
is entitled to subspecific rank. In the tributaries of the lake, the typical form is
always present.

In western New York, it has been reported from the St. Lawrence-drainage as

ORTMANN: monograph of the naiades of PENNSYLVANIA.

101

well as from the Erie Canal and the Mohawk River (Call) and it belongs to the
few species, which have gone along this route probably in very recent times.
Whether this stock came from Lake Erie or from the upper Allegheny, remains
to be seen.

Elliptio dilatatus sterkii Grier (1918).

Unio gibbosus Barnes (Sterki, 1907a, p. 392) small form; Unio gibbosus Barnes
(Ortmann, 19096, p. 203) lake form; Unio gibbosus Barnes (Walker, 1913,
p. 22, map, fig. 2 on p. 30) ; Elliptio dilatatus sterkii Grier, 1918, p. 9.

Plate VIII, fig. 3.

Records from Pennsylvania :

Ortmann, 19096, p. 203 (as form of Unio gibbosus from Lake Erie).

Grier, 1918, p. 9.

Characters of variety: Distinguished from typical E. dilatatus by small size,
rather swollen shell, (with the diameter over thirty percent, while it is less in the
typical form), the more anterior position of the beaks, and lighter color of the
epidermis, which is from yellowish olive to brown (sometimes dark brown), with
more distinct rays (chiefly when young). Generally in old shells the posterior
slope or the posterior end of the shell is lighter in color (yellowish to rusty brown) •
The growth-rests are rather regular, and rather well-marked by concentric dark
bands. Nacre mostly lighter, white to light purple, but sometimes as dark as
in the typical form.

L.

H.

D.

Pr.ct.

1.

Erie,

Cat. No. 61.3228 (largest)

.... 87 mm.

45 mm.

33 mm.

.38

2.

Erie,

Cat. No. 61.4628 (d^) type set. . . .

.'...86 “

40 “

33 “

.38

3.

do.

do.

(9)

do

....77 “

37 “

27 “

.35

4.

do.

do.

(9)

do

79 it

33 ‘

23 “

.32

5.

do.

do.

(c7)

do

....58 “

29 “

18 “

.31

Soft parts: They agree entirely with those of typical dilatatus. Glochidia
not known.

Breeding season: Gravid females (with eggs) have been observed on July 8
and 12, 1910.

Remarks: With a rather large number of specimens (over fifty) before me, I
am convinced that this is, as Grier has pointed out, a good local race of E. dilatatus.
Although the normal form locally does not grow very large, the lake-form is always
more swollen (see measurements) , and this gives to the shell a rather subcylindrical
shape, chiefly so in its anterior part. In the color of the epidermis, young speci-
mens of sterkii are distinctly lighter than normal dilatatus, and in consequence of

102

MEMOIRS OF THE CARNEGIE MUSEUM.

this, the rays are better visible, and the growth-rests become more marked. When
older the epidermis becomes darker, and may be even blackish, but in many old
specimens it remains light, chiefly so at or near the posterior end. This part of
the shell is often covered with a growth of other organisms. The regularity of the
growth-rests is rather variable, but in well-developed specimens it is quite striking.
In shape young specimens are more like normal dilatatus. In fact sometimes they
are indistinguishable except by the color.

Localities represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania (type locality).

Lake Erie, off North shore of Presque Isle, Erie, Erie Co., Pennsylvania (depth 10 feet).

Lake Erie, Cedar Point, Erie Co., Ohio (C. Brookover); La Plaisance Bay, Monroe Co., Michigan (C.

Goodrich); Port Colborne, Welland Co., Ontario, Canada (C. Goodrich).

Distribution and Ecology (See fig. 10 and Walker, 1913, fig. 2, p. 30) : Type
locality: Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania. Type set:
Carnegie Museum Cat. No. 61.4628 (not 4268, as Grier states!).

This form is exclusively known from Lake Erie. According to my observa-
tions it lives in Presque Isle Bay, preferably along the North shore of the bay, in
the characteristic fine sand, in from one to five feet of water, and chiefly at the
edge or within the Juncus americanus formation (among “rushes”). It also is
frequently found among Cliara patches. However, I also found it on the south
shore of the bay in gravel and shingle, and one specimen was obtained by the
“sand-sucker” in the open lake, in about ten feet of water.

Sterki (1907a, p. 392) and Walker (1913, p. 22) are the only authors, who
previously have mentioned this form from Lake Erie, but without separating it
from the main form. Grier (1918) first recognized it as a variety.

Note: It is interesting to compare specimens obtained under similar ecological
conditions at other localities. A very remarkable form is E. dilatatus from Chaut-
auqua Lake, Chautauqua Co., New York. I have eighteen specimens collected
by various parties at Bemus Point and Celeron. This is a form distinctly inclining
towards the var. sterkii. It is rather small (longest 79 mm.), is also slightly more
swollen than the true dilatatus, and has the beaks a little more anterior; but with
regard to color, the Chautauqua form does not differ from dilatatus. This is, of
course, no intergrade (genetically), but has developed independently, coming up
into Lake Chautauqua from the upper Allegheny.

In Conneaut Lake, Crawford Co., Pennsylvania, a form of E. dilatatus exists,
which is not at all different from the normal one, except that it is rather small.
It is rare in this lake, but more abundant in the outlet, and assumes, in French
Creek, to which the outlet flows, the typical size.

OETMANN: monograph of the naiades of PENNSYLVANIA.

103

The only tributary of Lake Erie in Pennsylvania, Conneaut Creek, contains a
small race of typical E. dilatatus. There are no indications, that this has come
up from the lake. It may as well have crossed over from the Beaver or French
Creek drainages.

Of course E. dilatatus sterkii must have come into the lake from the Ohio
system, and the Wabash-Maumee route is the first to be considered (See Walker,
1913), but it may have crossed elsewhere, since the typical form ascends in many
places far into the headwaters. The remarkable thing is that in all tributaries of
the lake the normal dilatatus is present, in Michigan at Ann Arbor), Marshall,
1895) in Ohio in the Cuyahoga (Dean, 1890) in Indiana in the drainage of the
Maumee, St. Marys, and St. Josephs Rivers (Call, 1900). Call mentions the fact
that there is in the lakes of northern Indiana, a form which is flatter and thinner.
This, of course, would not lead to sterkii.

Elliptic violaceus (Spengler) (1793).

Unio violaceus Spengler. Haas, 1913, p. 54, text-fig. 2; Unio complanatus
(Dillwyn) Simpson, 1914, p. 651.

Plate VIII, figs. 4, 5.

Records from Pennsylvania:

Say, 1817 (Delaware and Schuylkill Rivers) as purpureus (See Binney, 1858).

Haldeman, 1844 (Lancaster Co.).

Lea, Obs. IV, 1848 (Cobbs Creek, La Grange, near Philadelphia) (fuligmosus).^^

Gabb, 1861 (League Island and Schuylkill River, Philadelphia).

Bruckhart, 1869 (Lancaster Co.).

Hartman & Michener, 1874 (Chester Co.).

Schick, 1895 (Delaware and Schuylkill Rivers, Corinthian Reservoir, Philadelphia; Canal at Manayunk,
Philadelphia Co.; Munckinipattus Creek, Glenolden, Delaware Co.).

Ortmann, 19095, p. 208.

Caffrey, 1911 (Lehigh and Delaware Rivers, Northampton Co.).

Characters of the shell: Shell moderately large and moderately heavy. Outline
subtrapezoidal, more or less elongate, generally almost twice as long as high, or
even longer. Anterior end rounded, posterior end slightly produced and slightly
pointed; lower margin more or less convex, often nearly straight. Upper margin
subparallel to the lower margin, forming an angle with the obliquely descending
posterior margin. Beaks not prominent, at variable distance from the anterior
end, but not close to it. Beak-sculpture distinct, consisting of five to six ridges,
the first two or three curved and subconcentric, those following running in the

Cobbs Creek is a branch of Darby Creek, near Essington, Philadelphia County.

104

MEMOIRS OF THE CARNEGIE MUSEUM.

direction of the growth-lines, being subparallel and nearly straight in the middle,
and curving up in front and behind, the posterior curve being angular, with a
slight swelling (See Marshall, 1890, fig. 6). Shell not, or very little, swollen, gen-
erally more or less compressed, chiefly so on the posterior slope, with a blunt, more
or less distinct posterior ridge. Sides of the disk generally flat. Surface without
sculpture.

Epidermis yellowish, to brown and blackish. In young specimens, more or
less distinct greenish rays are present. The epidermis rarely is lighter and greenish
in color, with more distinct rays. The latter are capillary or somewhat broader,
straight. In older shells, all traces of rays generally disappear. Growth-rests
more or less strongly marked, but often (piite indistinct.

Hinge-teeth well-developed, but not very heavy. Pseudocardinals divergent,
ragged. Interdentum practically absent. Lateral teeth long, gently curved.
Beak-cavity very shallow. Dorsal muscle-scars in the beak-cavity. Nacre white
or colored, with an immense range of variation, through all shades of salmon, pink,
red, purple, and blueish, with coppery or bronze lustre or iridescence; often whitish
with lurid tints (greenish and grayish, as if discolored).

No sexual differences in the shell.

L. H. D.

Size: 1. Flinton, Cat. No. ()1.3797 (largest from

Pennsylvania)

120 mm.

65 mm.

32

mm.

2. Tioga, Cat. No. ()1.4312

no “

57 “

25

((

3. Selinsgrove,yCat. No. 61.4643

100 “

.58 “

35

(i

4. Manavunk, Cat. No. 61.1832

88 “

47 “

27

Sojt parts (See Ortmann, 1912, p. 269). Glochidia: ibid., PL 19, fig. 1.

Breeding season: For gravid females the following dates are at hand: April
26, 1909; May 3, 1909; May 4, 1909; May 6, 1912; May 9, 1911; May 10, 1912;
May 11, 1912; June 3, 1912; June 4, 1912; June 5, 1912; June 7, 1912; June 10,
1912; June 12, 1912; June 13, 1912; June 14, 1910; July 9, 1914; July 11, 1914;
July 16, 1908. Glochidia have been found as early as June 7, and discharging
females have been observed on June 7 and 13, and July 9 and 11. According to
these dates this is a tachytictic form, breeding from the end of April to July, but
the season may extend to August, as Conner (1907, p. 88) states. The discharging
females expelled their placentae whole into the surrounding water.

Remarks: This species is quite characteristic and easily recognized, in spite
of its immense range of variation. The trapezoidal outline (with the upper and
liosterior margins forming an angle), and the flat sides of the shell are reliable
features, and in most cases the shell is distinctly compressed. If more swollen.

OHTMANN: monograph of the naiades of PENNSYLVANIA.

105

the greatest width is situated in the region of the jiosterior ridge, but not in the
anterior part of the shell, as in E. dilatatus. This anterior swelling, and the tapering
posterior end of the shell, distinguish this latter species from E. violaceus.

In other respects E. violaceus is extremely variable. First of all in size and
shape (longer or shorter), then in the degree of compression (compare our measure-
ments). Furthermore the color of the outside and of the nacre is variable, and
finally the epidermis may be more or less smooth, or may be roughened by con-
centric lamellse. Since the shell is not very particular as to station (see below),
it is not astonishing that the responses to the environment are numerous, and
that it assumes in many places peculiar features, which, however, are not at all
constant, and may turn up anywhere under proper conditions, and are connected
by innumerable intergrades. There is no tendency in Pennsylvania to develo])
local races or varieties, and I am perfectly satisfied that there is only one species
in our region.

‘‘Species-making” within this form has gone beyond all bounds, and in a
large number of Lea’s “species,” chiefly from the southern states, the question
may be raised whether Lea was actually in earnest, when proposing them, or
whether he only wanted to mystify contemporaneous and subsequent students
of naiadology. Great credit is due to Simpson for straightening out the worst of
this tangle; but I think that Simpson has not gone far enough. I shall mention
here a few additional instances of species created by Lea, which in my opinion
simply fall as synonyms under E. violaceus, merely representing individual phases.
This chiefly concerns such forms as are actually found in our state.

Unio complanatus jejunus (Lea): Simpson (1914, p. 658) imiies jeju7ius and
percoarctatus Lea, and makes this a variety of the present species. Both are
southern types (from the Carolinas), and both are said to be more compressed,
while jejunus is reported to have an inclination to be biangulate behind, and
percoarctatus is said to have the surface covered with loose concentric strise. All
these characters are found in the Pennsylvanian E. violaceus in certain individuals.
They may be very much compressed, they have very often a tendency to be bi-
angulate behind, and they have often a rough epidermis, with loose strise. I have
specimens, which completely agree with jejunus, as well as with percoarctatus, and
sometimes such specimens (See Plate VIII, fig. 5) prevail at certain localities,
but they are always connected with the normal form by intergrades.

Unio roanokensis Lea, and northamptonensis Lea: Simpson (1914, p. 666)
regards these as one species, distinct from our species. The form northamptonensis
is reported from as far north as Massachusetts, and might be expected in Penn-

106

MEMOIRS OF THE CARNEGIE MUSEUM.

sylvania. Both shells are described as being large, flat, biangulate behind, while
northamptonensis is said (Simpson, 1900, p, 728 footnote) to have in addition the
posterior point elevated above the base line. Such shells may be found anywhere
in Pennsylvania. Specimens from Hartford, Connecticut, one of the type localities
of northamptonensis, and which are in the Carnegie Museum, are in fact normal,
large representatives of E. violaceus.

Unio fuliginosus Lea. This shell from Cobbs Creek, near Philadelphia, is
quoted twice by m}^son in 1900, first (p. 722) as synonym of complanatus, then

• Elliptis violaceus.

■ Elliptis cupreus.

+ Elliptis fisherianus.

'(p. 727) as of icterinus Conrad; but in 1914 (p. 665) only the latter reference is
given. I do not know icterinus, which is southern. The fuliginosus Lea, however,
does not agree with the original description of icterinus (Conrad, Mon. 4, 1836,
PL 18, fig. 2), and I think, the first reference of Simpson is correct, and the Penn-
sylvanian fuliginosus is identical with E. violaceus.

Localities in Pennsylvania represented in the Carnegie Museum:

Delawar e-drainage:

Delaware River, Penns Manor and Yardley, Bucks Co.; Shawnee, Monroe Co.

White Clay Creek, Avondale, Chester Co.

Schuylkill Canal, Manayunk, Philadelphia Co.

Wissahickon Creek, Flourtown, Montgomery Co. (P. A. Keppelmann).

Little Neshaminy Creek, Grenoble, Bucks Co.

Common Creek, Tullytown, Bucks Co.

Princess Creek, Kunkletown, Monroe Co.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

107

Meniolagomeka Creek, Smith Gap, Monroe Co.

Lizard Creek, Mantz and West Penn, Schuylkill Co.

Mahoning Creek, Lehighton, Carbon Co.

Susquehanna-drainage :

Susquehanna River, York Furnace and York Haven, York Co.; Duncannon, Perry Co.; Selinsgrove,
Snyder Co.

Codorus Creek, York, York Co.

Conewago Creek, York Haven, York Co.; Table Rock, Adams Co.

Conodoguinet Creek, Carlisle, Cumberland Co.

Shermans Creek, Duncannon, Perry Co.

Juniata River, Juniata Bridge, Perry Co.; Lewistown, Mifflin Co.

Canal, Mifflintown, Juniata Co. (D. A. Atkinson).

Lost Creek, Mifflintown, Juniata Co. (D. A. Atkinson).

Cocolamus Creek, Cocolamus, Juniata Co. (D. A. Atkinson).

Raystown Branch Juniata River, Ardenheim, Huntingdon Co.; Everett and Mount Dallas, Bedford Co.;
Lutzville, Bedford Co. (J. F. L. Raschen); Dunning Creek, Bedford, Bedford Co. (J. F. L. Raschen);
Shobers Run, Bedford Springs, Bedford Co. (A. Koenig).

Frankstown Branch Juniata River, Huntingdon and Alexandria, Huntingdon Co. (D. A. Atkinson);
Hollidaysburg, Blair Co.

West Branch Mahantango Creek, Richfield, Juniata Co. (D. A. Atkinson).

Middle Creek, Freeburg, Snyder Co. (D. A. Atkinson).

Penns Creek, Selinsgrove, Snyder Co. ‘ •

Canal, Watsontown, Northumberland Co. (D. A. Atkinson).

West Branch Susquehanna River, Williamsport, Lycoming Co. (D. A. Atkinson).

Bald Eagle Creek, Milesburg, Center Co.

Driftwood Branch Sinnamahoning Creek, Driftwood, Cameron Co.

Beaver Dam Creek, Flinton, Cambria Co. (D. A. Atkinson).

Chest Creek, Patton, Cambria Co.

Cush Cushion Creek, Green Township, Indiana Co. (D. A. Atkinson).

North Branch Susquehanna River, Tunkhannock, Wyoming Co.

Chemung River, South Waverly, Bradford Co.

Millrace of Crooked Creek, Tioga, Tioga Co.

Potoviac-drainage :

East Branch Little Antietam Creek, Waynesboro, Franklin Co.

Conococheague Creek, Greencastle and Scotland, Franklin Co.

West Branch Conococheague Creek, Mercersburg Junction, Franklin Co.

Great Tonoloway Creek, Thompson Township, Fulton Co.

Localities in Pennsylvania represented in the Philadelphia Academy of Natural
Sciences:

Sacony Creek, Kutztown, Berks Co. (H. K. Deisher).

Big Neshaminy Creek, “ Edderton,” Bucks Co. (H. W. Fowler).

“ Guinea Creek,” Woodbourne, Bucks Co. (H. W. Fowler).

Probably Eddington.

No creek of this name occurs on the map of the U. S. Topographic Survey (Sheet Burlington).
The creek at Woodbourne is a branch of Mill Creek.

108

MEMOIRS OF THE CARNEGIE MUSEUM.

Buslikill Creek, Belfast, Northampton Co. (H. W. Fowler).

Susquehanna River, York Furnace, York Co. (W. Stone).

Little Swatara Creek, Jonestown, Lebanon Co. (W. H. Zehring) (C. H. Conner).

Juniata River, Newton-Hamilton, Mifflin Co. (H. T. Mather, Jr.).

Localities on the New Jersey side of the Delaware in the Philadelphia Academy
of Natural Sciences:

Delaware River, Washington Park, near Newbold, Gloucester Co. (C. H. Conner); Delanco, Burlington
Co. (H. A. Pilsbry); Burlington Island, Burlington Co. (H. W. Fowler).

Other localities represented in the Carnegie Museum:

St. Laivrence-drainage:

Spider Bay of Georgian Ba}'', Sans Souci, Parry Sound Co., Ontario, Canada (H. Kahl).

Muskoka Lake, Bala, Muskoka Co., Ontario, Canada (A. S. Daggette).

Severn River, Gloucester Pool, Muskoka Co., Ontario, Canada (0. A. Peterson).

Pigeon Lake, Bobcaygeon, Victoria Co., Ontario, Canada (G. H. Clapp).

Sandy Lake, Peterboro Co., Ontario, Canada (G. H. Clapp).

St. Lawrence River, Montreal, Quebec, Canada (Hartman collection); Bluff Island and Grindstone
Island, near Clayton, Jefferson Co., New York (Miss A. H. Robinson) (H. Kahl).

Lake Ontario, Hamilton Bay, Wentworth Co., Ontario, Canada (Miss R. Buchert); Braddock Bay,
Manitou Beach, Monroe Co., and Sodus Point, Wajme Co., New York (R. H. Santens).

Genesee River, Roche.ster, Vlonroe Co., New York (R. H. Santens).

Lake Champlain, New York (Hartman collection).

Nexv England:

Fish River, Eagle Lake, Square Lake, and Cross Lake, Aroostook Co., Maine (0. 0. Nylander).

Little Madawaska River, Westmorelaiul, Aroostook Co., Maine (0. O. Nylander).

Piscataquis Co., Maine (C. Goodrich, donor).

Otisheld, Cumberland Co., Maine (G. H. Clapp, donor).

Lilly Lake, Pittsfield, Merrimack Co., New Hampshire (Mrs. L. D. Thompson, donor).

Shawsheen River, Bedford, Middlesex Co., Massachusetts (Mrs. L. D. Thompson).

Haughtons Pond, Blue Hills, Norfolk Co., Massachusetts (Mrs. L. D. Thompson, donor).

Connecticut River, Hartford, Hartford Co., Connecticut (Hartman collection).

New York and New Jersey:

North Branch Susquehanna River, Binghamton, Broome Co., New York (H. H. Smith).

Culvers Pond, Sussex Co., New Jersey (J. F. L. Raschen).

Stony Brook and Delaware-Raritan Canal, Princeton, Mercer Co., New Jersey.

Delaware River, Fish House, Camden Co., New Jersey.

Only those are given for which I either can vouch personally, or which are from reliable sources.
The Carnegie Museum possesses many more, chiefly from the southern states, which may be correct or
wrong, and which are referred to various species of Lea, which cannot properly be identified at present.
Specimens from Catawba River, Bridgewater, Burke Co., North Carolina, collected by myself, correspond
well with U. percoarctatus Lea, which is from the same river in Gaston Co. a little lower down. No
typical E. violaceus was met with here, but associated with it is a form appearing quite distinct, but
which I have been unable to identify. Walker, to whom I have sent specimens, did not wish to express
an opinion in regard to them.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

109

Barton Run, below Tomlisons Mill (= Jennings Mill), Marlton, Burlington Co., New Jersey (C. 11.
Conner).'^'’

Potomac-drainage:

Potomac River, Cabin John, Montgomery Co., Maryland (J. D. Haseman); Hancock, Washington Co.,
Maryland; Cherry Run, Morgan Co., West Virginia.

Wills Creek, Ellerslie, Allegheny Co., Maryland.

South Branch Potomac River, Southbranch and Romney, Hampshire Co., West Virginia.

Shenandoah River, Harpers Ferry, Jefferson Co., West Virginia.

North Fork Shenandoah River, Broadway, Rockingham Co., Virginia.

South Fork Shenandoah River, Elkton, Rockingham Co., Virginia.

South River, Waynesboro, Augusta Co., Virginia.

Southern localities:

Rappahannock River and Marsh Run, Remington, Fauquier Co., Virginia.

Mountain Run, Culpeper, Culpeper Co., Virginia.

Rapidan River, Rapidan, Culpeper Co., Virginia.

North River, Lexington, Rockbridge Co., Virginia.

Calf Pasture River, Goshen, Rockbridge Co., Virginia.

Jackson River, Covington, Allegheny Co., Virginia.

Roanoke River, Salem, Roanoke Co., Virginia.

Tinker Creek, Roanoke, Roanoke Co., Virginia.

Mason Creek, Salem, Roanoke Co., Virginia.

Distribution and Ecology in Pennsylvania (See fig. 11): This is the commonest
form of Naiades in the Atlantic-drainage in Pennsylvania, and is as characteristic
and as abundant on the eastern side of the Alleghenian divide, as is E. dilatatus
on the western, in fact, it is even more universally distributed than the latter.
(Ortmann, 1913a, pp. 361, 362). It apparently has no ecological preferences,
being found practically in any permanent bo,dy of water; in canals and reservoirs
with quiet water and muddy bottom, as well as in large rivers with strong current
and heavy gravel and rocks. In the small creeks it goes up very far into the head-
waters, and is found, for instance, in the small tributaries of the West Branch of
the Susquehanna, close to the divide as in Cush Cushion Creek in Indiana County.
Here it is only about twenty miles away from the nearest locality of E. dilatatus
in the Allegheny-drainage in Indiana County. (Yellow and Little Mahoning
Creeks.) Also in Virginia, similar observations have been made (See Ortmann,
1913a). In the region, where the headwaters of New River and those of the
James and Roanoke Rivers come together, the two species are found immediately
west and east of the divide.

The ability of E. violaceus to live everywhere under a great variety of environ-
mental conditions undoubtedly accounts for its great variability, as well as its

Quite a peculiar form, unusually swollen, with white nacre, which requires further investigation.

no

MEMOIRS OF THE CARNEGIE MUSEUM.

tendency to develop many different phases, which may turn up anywhere under
proper conditions, but which do not lead to the development of geographical races,
at least in our territory.

General distribution: Type locality, North America (Spengler).

According to Simpson (1900, p. 725), E. violaceus is found on the Atlantic
side from the St. Lawrence to Georgia. The southern boundary is extremely un-
certain, and is obscured by a great development of local forms, the standing of
which is very doubtful. Northward, it goes to Maine (Jackson, 1908, Lermond,
1909, Nylander, 1914), and, according to Call, to Nova Scotia. From the lower
St. Lawrence-drainage, it has been reported from many localities in Vermont,
New York, and Canada, up to and into Lake Ontario. It has never been found
in Lake Erie, but turns up again in the upper lakes district, in western Ontario
(see our localities in Sandy and Pigeon Lakes, and in Muskoka and Parry Sound
Cos.). It has been discovered in various places in northern Michigan, as far as
Lake Superior (See Walker, 1898, p. 5; 1913, map on p. 30, and Winslow, 1917,
p. 11). An isolated lofcality is the one reported by Sterki (1907a, p. 393) from a
mill-race at New Philadelphia, Tuscarawas Co., Ohio, and recently I found a single
specimen in Grand River, Eagleville, Ashtabula Co., Ohio.

This westward extension of the range apparently was not by way of Lake
Erie, but from the lower St. Lawrence, in the region about Ottawa, in the direct
line toward Lake Huron, so that in Michigan it is a recent immigrant from the
North. Walker (1913) has treated this question in detail: the migration was by
way of the Trent or Nipissing route {1. c., pp. 44, 45, fig. 4).

The established range of this species is unique, and larger than that of any
other form of the Atlantic slope. Most peculiar is the westward extension in the
north. There ‘is only one species, which might be compared with this, Eurynia
nasuta, but the range of this from west to east goes by way of Lake Erie, and is
undoubtedly governed by different laws, as I have shown (Ortmann, 1913a, pp.
378 ff.).

Elliptio cupreus (Rafinesque) (1820).

Unio productus Conrad. Simpson, 1914, p. 690.'^^

Plate VIII, fig. 6.

Records from Pennsylvania:

Ortmann, 1910, p. 117 (Fulton Co.).

Conrad (1834, p. 8) says that the shells of the type of U. purpureus (= E. violaceus) are in the
Savannah, Oconee, and Ocmulgee Rivers, and also in Flint River, Georgia, but that to the west of this
he did not find them.

U. fisherianus Reeve (1865, PI. 24, fig. 113) is not U. fisherianus of Lea, but clearly the present

species.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

Ill

Characters of the shell: Shell rather small and not heavy. Outline elongated
trapezoidal, or elongated ovate, lance-head shaped, a little over twice as long as
high. Greatest height of the shell about in the middle at the ligament (mostly
height of shell about the same from the beaks to the end of the ligament) . Anterior
end rounded, posterior end tapering and bluntly pointed. Lower margin more or
less curved, but often almost straight in the middle. Upper margin straight or
somewhat curved, nearly parallel to the lower margin, and forming a more or less
distinct angle with the upper posterior margin. Beaks not prominent, placed
rather anteriorly, but at a certain distance from the anterior end. Beak-sculpture
identical with that of E. violaceus, but sometimes the straight portion of the bars,
in their middle part, is slightly and indistinctly sinuated. Shell not swollen, more
or less compressed, with the sides of the disk rather flat. A rounded, indistinct
posterior ridge is present. Surface without sculpture.

Epidermis greenish olive, brownish olive, yellowish brown to brown, or even
blackish. Young specimens sometimes with rather indistinct, greenish rays, of
the same character as those of E. violaceus, but in most cases no traces of rays
are visible. Growth-rests generally not marked off by darker color.

Hinge-teeth well-developed, but not very strong, of the general character of
those of E. violaceus. Beak-cavity shallow. Dorsal muscle-scars in the beak-
cavity. Nacre whitish, lurid, or of some shade of red or purple, lighter or darker.

often with coppery iridescence.

No sexual differences in the shell.

L. H. D.

Size: 1. Romney, Cat. No. 61.5898 (extralimital) 88 mm. 38 mm. 23 mm.

2. Fulton Co., Cat. No. 61.4323 (largest from Pa.) . . 70 “ 30 “ 15 “

3. do. do. 66 “ 30 “ 16 “

4. do. do. 60 “ 27 “ 16 “

5. do. do. 49 “ 23 “ 11 “

Soft parts (See Ortmann, 1912, p. 270). Gravid females have subsequently
been found, and the structure of the marsupium agrees fully with that of E. violaceus.
Glochidia: L. 0.20, H. 0.20 mm., similar in shape to those of E. violaceus (which
are 0.20 X 0.19 mm.).

Breeding season: Gravid females were found on Alay 6, 1912; June 3, 1912;
June 4, 1912; June 7, 1912; June 8, 1912. All these were from the Potomac,
Rappahannock, and James drainages, outside of Pennsylvania. At the earliest
date, glochidia in some individuals were already present.

Remarks: Obliquaria cuprea Rafinesque (1820, p. 304) is this species, as is
evident from the description, the very poor (exaggerated) figure, and the comparison

112

MEMOIRS OF THE CARNEGIE MUSEUM.

with U. nasuta Say. Rafinesque gives the Monongahela and Potomac Rivers as
habitats, but the first locality apparently is an error. (On p. 294 Rafinesque de-
scribes a Unto fasciata cuprea, but this does not conflict with Ohliquaria cuprea).

This species is allied to the type of E. violaceus, but differs in its extreme
elongation and lanceolate shape. The subtrapezoidal outline is obscured, but at
least in young specimens is indicated by the presence of an angle between the
upper and the posterior margins. It is hardly possible to confuse this species
with E. violaceus, but it has a much closer resemblance to the following {E. fisheri-
anus), and has been mixed up with this by various writers. It is possible that
E. fisherianus is merely a variety of E. cupreus. There are other '‘species” in the
southern states, which group with this form, but they require further study.

This shell (as well as E. fisherianus) has an outward resemblance to Eurynia
nasuta (Say), and was supposed to be nearly allied to it by the earlier authors,
but the anatomical structure is entirely different.

Localities represented in the Carnegie Museum:

Great Tonoloway Creek, Thompson Township, Fulton Co., Pennsylvania.

Potomac River, Hancock, Washington Co., Maryland.

South Branch Potomac River, Southbranch and Romney, Hampshire Co., West Virginia.

Rappahannock River, Virginia (G. H. Clapp, donor).

Marsh Run, Remington, Fauquier Co., Virginia.

Mountain Run, Culpeper, Culpeper Co., Virginia.

North River, Buena Vista and Lexington, Rockbridge Co., Virginia.

Calf Pasture River, Goshen, Rockbridge Co., Virginia.

Jackson River, Covington, Alleghany Co., Virginia.

Locality represented in the Philadelphia Academy of Natural Sciences:

Shenandoah River, Harpers Ferry, Jefferson Co., West Virginia (G. W. Tryon, Jr.).

Distribution and Ecology (See fig. 11): Type-locality, Potomac River '"(Rafin-
esque) .

Conrad’s Unio productus, which undoubtedly is this species, is given from
the Savannah River, Augusta, Richmond Co., Georgia. But Simpson (1900, p.
735) mentions only North Carolina, Virginia, and Maryland. But later (1914,
p. 691) he adds the type-locality. The southward range of this species as yet
remains obscure, and furthermore very few exact localities are on record. Besides
those given by myself, I know of only one which is reliable, that furnished by
Pilsbry (1894) from Sideling Creek, Allegany Co., Maryland.

As I have suggested elsewhere (Ortmann, 1913a, p. 320), Conrad’s Unio lanceolatus (1846) from
the Upper James-drainage, is probably this species.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

113

I have located this species in the drainages of the Potomac, Rappahannock,
and James, in Maryland, Virginia, and eastern West Virginia, and also in a northern
tributary of the Potomac (Great Tonoloway Creek) in southern Pennsylvania,
just North of the Mason and Dixon line. I have hunted for this species in other
tributaries of the Potomac in Pennsylvania, in Antietam and Conococheague
Creeks, but failed to find the slightest trace of it.

Where I collected this form, it is not at all rare. I found it either in gravel,
or among larger rocks, in mud-filled interstices, but generally not in strong current,
but in quiet coves and eddies. It also seems to prefer the smaller streams to the
larger rivers.

With the exception of the Rappahannock, I never found it East of the Blue
Ridge, on the Piedmont Plateau or Coastal Plain, but it should be expected to be
more frequent there. The relation of its distribution to that of the next species
{E. fisherianus) should be ascertained.

Elliptic fisherianus (Lea) (1838).

Unio fisherianus Lea. Simpson, 1914, p. 692.

Plate VIII, fig. 7.

Records from Pennsylvania:

Gabb, 1861 (Schuylkill River, above Girard Bridge, Philadelphia).

? Hartman & Michener, 1874 (White Clay Creek, Chester Co.) (see below).

Characters of the shell: Very close to E. cupreus, and differing chiefly by some-
what larger size, slightly more elongated shape, and greater taper in the posterior
part of the shell. The greatest height of the shell is situated more anteriorly, at
about the beaks, and from this part the shell decreases in height to the posterior
end. At the same time the posterior end is more elevated, so that the posterior
ridge is not straight, but curves up slightly behind, and the lower margin also is
more curved. In all other characters this form agrees with E. cupreus. The
epidermis is more greenish blackish, the nacre white to purple.

L.

H.

D.

Size: 1.

Kent Co., Del.

, Cat. No. 61.4645

106 mm.

40 mm.

26 mm.

2.

do.

do.

88 “

35 “

23 “

3.

do.

do.

73 “

27 “

19 “

4.

do.

do.

57 “

22 “

14 “

Soft parts, glochidia, and breeding season unknown.

Remarks: According to the material at hand, this species stands very close
to E. cupreus, and it possibly is only a local race (lowland form) of it. However

114

MEMOIRS OF THE CARNEGIE MUSEUM.

intergrades have not been observed, but may exist in the lower parts of the rivers.
Attention should be paid to this question.

Localities represented in the Carnegie Museum:

Mill-pond of Ratcliff Creek, Chestertown, Kent Co., Maryland (E. G. Vanatta).

Choptank Mills, Kent Co., Delaware (S. N. Rhoads).

Localities represented in the Philadelphia Academy of Natural Sciences:

Church Hill, Queen Anne Co., Maryland (E. G. Vanatta).

Potomac River, Washington, D. C. (G. W. Tryon, Jr.) (John Ford).

Distribution and Ecology (See fig. 11): Type locality, Head of Chester River,
Maryland (Lea).

Simpson reports this species from: “Virginia, IMaryland, Pennsylvania,” and
Rhoads (1904) from two localities in Delaware. There is no doubt that “Penn-
sylvania” is founded iqion the records mentioned above (Gabb, and Hartman &
Alichener). Gabb reports only a single specimen from his locality, the lower
Schuylkill River, and since this evidently offers the necessary conditions, there
is no reason to doubt the fact that the sjiecies has occurred there. However, I
entertain strong doubts that this species ever existed in White Clay Creek in Penn-
sylvania. I have been at White Clay Creek, at Avondale, Chester Co., Pennsyl-
vania, and here it is an upland stream, and does not at all present the conditions,
which I regard as being favorable for this species. It is possible that Hartman &
Michener found this species farther down in this creek, in the lowlands of the state
of Delaware, but not in Pennsylvania.'^^

The evidence for the existence of this species in Pennsylvania rests upon the
single individual reported by Gabb. It is not impossible that it may be found
again, and the chance to rediscover it will be in or near the estuary of the Dela-
ware, for according to the records at hand this species seems to prefer the Coastal
Plain, living in slowly running rivers, ponds, or similar stations; it is a lowland
form, as I have pointed out elsewhere (Ortmann, 1913a, p. 361).

Subfamily ANODONTINTi (Swainson) Ortmann (1910).

Ortmann, 1910, p. 117; 19116, p. 336; 1912, p. 224.

Key to the Genera of the Anodontin^.

ai. Beak-sculpture double-looped, with a sharp reentering angle or a sinuation. Mantle connection
between anal and supra-anal openings moderate or very long. No tendency to unite the
inner lamina of the inner gills with abdominal sac.

In addition to the above locality: Seaford, Sussex Co., Delaware.

The Carnegie Museum acquired in the Hartman collection a single set of this species, but there
is no indication that it came froni our state.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

115

bi. Hinge-teeth more or less well-developed, at least the pseiidocardinals present. Shell generally

not very thin Lasmigona.

bi. Hinge-teeth completely absent. Shell thin, generally very thin Anodonta.

a 2. Beak-sculpture concentric. Mantle connection between anal and supra-anal openings moderate.

Sometimes a tendency to unite the inner lamina of the inner gills with the abdominal sac.

bi. Beak-sculpture fine. Hinge-teeth practically absent Anodontoides.

62- Beak-sculpture heavy, often very heavy. Hinge-teeth more or less developed or rudimentary.

Cl. Hinge complete, or at least the pseudocardinals present. Marsupium with simple ovisacs.

Alasrnidonta.

C2. Hinge-teeth rudimentary, but slight traces of pseudocardinals present. Marsupium with
the ovisacs subdivided into transverse compartments Slrophitus.

Genus Lasmigona Rafinesque (1831).

Symphynota Ortmann, 1912, p. 280; Symphynota, Simpson, 1914, p. 480; Las-
migona Frierson, 19146, p. 40.

Type Alasrnidonta costata Rafinesque.

Simpson has divided this genus into three subgenera, and Ortmann (1914,
p. 41) has added others, but the latter are extralimital. The nomenclature of the
genus and the subgenera has been revised in this paper in accordance with Frierson
(19146, p. 40) and Walker (1918).

Key to the Subgenera of Lasmigona.

fli. Lateral hinge-teeth present. Shell smooth. Beak-sculpture double-looped. Hermaphroditic.

Subgenus Platynaias.

tti. Lateral hinge-teeth obliterated. Shell smooth, or with ribs upon the posterior slope. Gonochoristic.
bi. Shell subrhomboidal or subelongate. Posterior slope with ribs. Beak-sculpture indistinctly

double-looped Subgenus Lasmigona.

bi. Shell broadly ovate-rhomboidal, not elongated, with high posterior wing. Posterior slope with
ribs, which may be obliterated. Beak-sculpture sharply double-looped. . .Subgenus Pterosyna.

Subgenus Platynaias Walker (1918).

Symphynota Simpson, 1914, p. 481; Ortmann, 1914, p. 42; Platynaias Walker,
1918, p. 2.

Type Unio viridis Rafinesque.

Two species belong to this subgenus, which may be distinguished as follows.

Key to the Species of the Subgenus Platynaias.

«!. Shell moderately large, subsolid, compressed. Outline more or less rhomboidal; western habitat.

L. (P.) viridis.

a 2. Shell small, thin, not so much compressed, Outline rather subovate; eastern habitat.

L. (P.) subviridis.

116

MEMOIRS OF THE CARNEGIE MUSEUM.

Lasmigona (Platynaias) viridis (Rafinesque) (1820).

Symphynota compressa Lea. Simpson, 1914, p. 481; Lasmigona viridis (Rafin-
esque) Frierson, 1915, p. 59.

Plate IX, figs. 1, 2.

Records from Pennsylvania:

Rhoads, 1899 (Beaver River, Wampum, Lawrence Co.).

Ortmann, 1909&, p. 196, 202.

Characters of the shell: Moderately large, subsolid, but not very heavy. Out-
line subrhomboidal, or subtrapezoidal, longer than high. Anterior end rounded,
posterior subtruncate. Lower margin slightly convex; upper margin straight,
more or less ascending and elevated posteriorly, forming a more or less distinct
wing at the upper posterior angle. In young shells the upper posterior margin is
symphynote. Beaks not prominent, placed in front of the middle of the shell.
Beak-sculpture distinct, consisting of about five bars, of which the first or the two
first are subconcentric, while those following are distinctly double-looped, with the
anterior loop rounded, the posterior angular, and slightly narrower than the an-
terior, a sharp re-entering sinus between them. Sometimes the last bars are some-
what irregular, interrupted or wavy in their anterior part, but this feature is vari-
able and often absent (compare Marshall’s figure, 1890, fig. 1). Shell rather flat
and compressed, disk only slightly convex. An indistinct, blunt posterior ridge
is present. Greatest diameter of the shell at, or immediately in front of, this
ridge. Behind the ridge the posterior slope appears more or less compressed and
winged. Surface without sculpture.

Epidermis yellowish to greenish, or, when old, blackish, with mqre or less
distinct green to blackish rays, which are straight, narrow or broad, often con-
sisting of bundles of capillary rays. Generally the posterior slope is darker green
on account of the more crowded rays. In old shells the rays become obscure, and
the whole surface turns greenish black. Growth-rests rather distinct, and mostly
marked by darker (brown) color.

Hinge-teeth well-develojied, but not heavy, rather delicate. Pseudocardinals
lamellate, directed forwards, and almost parallel to the hinge. Interdental pro-
jection of left valve well-developed, fitting into an interdental groove of the right
valve. Lateral teeth rather long, straight, well-developed, but thin. Beak-
cavity shallow. Dorsal muscle-scars in beak-cavity. Nacre white, often cream-
color or even pale salmon or flesh-color towards the beak-cavity.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

117

L. H. D.

Size: 1. Linesville,

Cat. No. 61.874

12 1 mm.

67 mm.

39

mm.

2. Shenango,

Cat. No. 61.4218

91 “

55

ii

29

ii

3. do.

do.

87 “

53

((

28

4. do.

do.

50 “

31

((

12

((

Soft parts (See Ortmaiin, 1912, p. 281). Glochidia (See Lea, Obs. VI, 1858,
PL 5, fig. 23; Ortmann, 1911, PL 89, fig. 10; Surber, 1912, PL 3, fig. 44). Siirber
gives the dimensions as 0.353 X 0.313 mm., while I found them to be: 0.34 X 0.28
mm.

This species normally is hermaphroditic, and I never have found a specimen
in Pennsylvania having the male structure of the gills. However on rare occasions
individuals with male structure seem to turn up. I have recorded (19115, p. 309)
such a case from Lake Erie, but this has remained the only one.

Breeding season: Gravid specimens were found on August 6, 1908; August
7, 1908; August 13, 1906; August 18, 1909; August 23, 1916; August 29, 1910;
Sept. 2, 1908; Sept. 3, 1908; Sept. 4, 1908; Sept. 7, 1908; Sept. 7, 1913; Sept.
10, 1906; Sept. 14, 1908; Sept. 18, 1916; Sept. 21, 1908; Sept. 27, 1909; Oct. 4,
1910; Oct. 10, 1907; Oct. 15, 1907; Oct. 19, 1908; and then again on May 14,
1908; May 22, 1908; May 23, 1908; June 2, 1908; June 17, 1909. The first
record for glochidia is on Sept. 7, while discharging specimens have been observed
on May 23, and June 2 and 17.

Thus this species is clearly bradytictic, beginning to breed early in August,
gravidity lasting till May and June of the following year, when the glochidia are
discharged. The interim between two succeeding breeding seasons falls into the
second half of June and in July.

Remarks: As to the specific name see the controversy between Frierson (1915,
p. 57) and Walker (1915, p. 74).

This is a species easily recognized by the shape and color of the shell, and by
the conformation of the hinge. It can only be confused with its nearest relative,
L. subviridis, but only in the young stage. Young specimens of L. viridis indeed
somewhat resemble older specimens of L. subviridis (Compare figs. 2 and 4, on
Plate IX) but they are always more distinctly trapezoidal, with well-developed
posterior wing. In other respects the size of full-grown specimens distinguishes
the two species at once. In Pennsylvania the geographical distribution is a good
diagnostic character, but this is not the case in New York, where the two species
overlap.

There is not much variation in the shell. The posterior wing may be more or
less developed, and the shell may be more or less elevated at the upper posterior

118

MEMOIRS OF THE CARNEGIE MUSEUM.

angle. The degree of compression is also variable; some specimens being more
swollen in the region of the posterior ridge than others. Furthermore, some speci-
mens are more elongated than others, but all these are individual variations, and I
never have noticed any distinct tendency to form local races.

Specimens from Conneaut Creek (Erie-drainage) do not differ from the normal
form of the Ohio-drainage. Specimens from the headwaters of the Genesee River
are identical with the type. The only two specimens I collected from Lake Erie
were not from the lake proper, but from beach-pools, cut off from the lake by

■ Lasmigona viridis.

• Lasmigona complanata.

-j- Lasmigona costata.

X Lasmigona costata eriganensis.

sand-bars. They are entirely normal, but have a peculiar rusty brown tint in the
epidermis, such as is found in other Naiades from Lake Erie, and especially in
specimens from the beach-pools. A third individual from the lake, from Cedar
Point, Ohio, is also normal in color.

Localities in Pennsylvania represented in the Carnegie Museum:

Little Beaver Creek, Cannelton (Miss Vera White) (H. H. Smith), Darlington and New Galilee, Beaver
Co.; Enon Valley, Lawrence Co.

Beaver-drainage :

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Brush Creek, Celia, Beaver Co.

Mahoning River, Mahoningtown, Lawrence Co.

Neshannock Creek, Eastbrook, Lawrence Co.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

119

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsvillc, Clarksville, Shenango, and
Jamestown, Mercer Co.; Linesville, Crawford Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

Little Shenango River, Greenville, Mercer Co. +

Randolph Run, Hartstown, Crawford Co.

A llegheny-drainage :

Cowanshannock Creek, Rural Valley, Armstrong Co. (N. VI. Grier).

Sandy Creek, Sandylake, VIercer Co.

French Creek, Utica, Venango Co.; Cochranton, Crawford Co.; Union City, Erie Co.

Conneaut Outlet, Conneautlake, Crawford Co.

Cussewago Creek, VIosiertown, Crawford Co. (H. & L. Ellsworth).

Conneauttee Creek, Edinboro, Erie Co.

Brokenstraw Creek, Garland, Warren Co.

Connewango Creek, Russell, Warren Co. ,

Potato Creek, Smethport, McKean Co. (P. E. Nordgren).

Lake-drainage:

Conneaut Creek, West Springfield, Erie Co.

Lake Erie, beach-pools of Presque Isle, Erie, Erie Co.

Genesee River, Genesee, Potter Co.

Cryder Creek, Genesee, Potter Co.

Other localities represented in the Carnegie Museum:

Lake Erie, Cedar Point, Erie Co., Ohio (C. Brookover).

Tenmile Creek, Lucas Co., Ohio (C. Goodrich) (lake-drainage).

Beaver Creek, Williams Co., Ohio (C. Goodrich).

Raisin River, Adrian, Lenawee Co., Michigan (C. Goodrich).

West Branch Nimishillen Creek, Canton, Stark Co., Ohio.

Tuscarawas River, Tuscarawas Co., Ohio (Holland collection).

North Fork Hughes River, Cornwallis, Ritchie Co., West Virginia.

Distribution and Ecology in Pennsylvania (See fig. 12) : The distribution of
this species in Pennsylvania is very peculiar, but, as we shall see later, there are
several similar cases. It belongs exclusively to the northwestern part of the state,
including Little Beaver Creek, the drainage of Beaver River and most of its tribu-
taries, some tributaries of the middle and upper Allegheny (Cowanshannock, Sandy,
French, Brokenstraw, Connewango, Potato Creeks), Lake Erie and its drainage,
and the upper Genesee River. It has never been found in the Allegheny proper
and its drainage from Armstrong Co. down, and is entirely missing in the Ohio
proper and in the whole Monongahela system.

Thus it seems as if this species in its distribution does not follow the courses
of the rivers, but rather goes across country in the northwestern part of the state,
and the relation of its area to that of the Glacial deposits is very close. Only in
Little Beaver Creek, below New Galilee, in Brush Creek (tributary to the Con-

120

MEMOIRS OF THE CARNEGIE MUSEUM.

noquenessing), Cowanshannock Creek (to middle Allegheny), and Potato Creek
(flowing into the uppermost Allegheny), is it outside of the Terminal Moraine,
but never far from it. The explanation of this peculiar fact will be attempted
elsewhere; here it suffices to direct attention to it, in order that the case may be
kept in mind.

As regards its ecology, L. viridis is in Pennsylvania distinctly a form of the
smaller streams, going up far into the headwaters, into streams which are indeed
the smallest in Pennsylvania, which contain mussels. Also Wilson & Clark
(1912a) have observed that it belongs to the small creeks in the upper Kankakee-
drainage in Indiana. Wherever I found this species, it seems to be averse to very
strongly flowing waters (in riffles) ; its usual station is near (below) riffles, where
there is a moderate flow of water over coarser or finer gravel, packed firmly by
fine sand or mud. It is generally not an abundant species, but locally it has been
taken in some numbers. In the beach-pools of Lake Erie I found it in the char-
acteristic sand of the lake-shores, with a scanty vegetation of alga3.

General distribiUiori: Type locality, rare in the Ohio, more common in the
Kentucky and small tributaries (Rafinesque).

Simpson (1900) condenses the range as follows: ‘‘Ohio and St. Lawrence
drainages areas; west to Arkansas, north through Nebraska to Wisconsin; Hudson
River.” This, however, does not bring out the chief feature of the distribution.
This species is distinctly more northern, and its main range is north of the Ohio in
Illinois, Indiana, Michigan, Ohio, and Pennsylvania; it crosses over into the lake-
drainage (Walker, 1913), and has in the St. Lawrence system a great extension
eastwards, crossing over in New York into the Atlantic-drainage (IMohawk and
Hudson Rivers). In Canada, it goes as far as the vicinity of Ottawa and Montreal
(Marshall, 1895), and extends to Vermont (Call, 1878, and Gray, 1883). It also
has been reported from Connecticut.^'^

In this eastern extension of its range it invades in part that of the next species
(L. subviridis), but it is not known whether it is found in any one locality associated
with the latter. Probably it went here (as did Fusconaia flava) by way of the
Erie Canal, although Call suggests the possibility of distribution by birds.

In the southward direction, in southwestern Pennsylvania, as indicated above,
and generally to the South of the Ohio River, it is missing or rare. I have never
found it in the Ohio proper from Pittsburg down to Cincinnati, although it is
mentioned in the Cincinnati list (Harper, 1896). I have found a single individual

New Haven Canal, according to Linsley (1845), but not mentioned by Perkins (1869). Adams
(1842) has distinguished this eastern form as var. 'plebeia (See Simpson, 1914, p. 483, and Johnson, 1915,
p. 25).

OETMANN: monograph of the naiades of PENNSYLVANIA.

121

in North Fork Hughes River, a tributary of the Tattle Ivanawha, and, for the present
I regard this as a stray specimen. Rafinesque gives it from Iventucky, also Baker
(1898a, .p. 60), but it is unknown from the Cumberland and Tennessee. The
distribution seems to be also very restricted in a southwesterly and westerly direc-
tion. The records from Arkansas and Iowa are very vague, and certainly need
verification (for Iowa, see Geiser, 1910) . It is absent from Call’s Arkansas list (1895) ,
and from Scammon’s Kansas list (1906). It is abundant in northern Illinois,
and passes into parts of Wisconsin and Minnesota, and also into eastern Nebraska.
But the exact boundaries in this region require further investigation.

Lasmigona (Platynaias) subviridis (Conrad) (1835).

Symphynota viridis Simpson, 1914, p. 484; Lasmigona subviridis (Conard) Frier-
son, 1915, p. 58.

Plate IX, figs. 3, 4.

Records from Pennsylvania :

Conrad, 1836 (Schuylkill River, Philadelphia; Lancaster; Juniata River).

Lea, 1838 (Juniata River, Hollidaysburg, Blair Co.).

Haldeman, 1844 (Lancaster Co.).

Conrad, 1856 (Schuylkill River, Phoenixville, Chester Co.; Delaware River, Morrisville, Bucks Co.:

“opposite Trenton”)-
Bruckhart, 1869 (Lancaster Co.).

Hartman & Michener, 1874 (Schuylkill River, Chester Co.).

Marshall, 1895 (Juniata River and Philadelphia).

Ortmann, 19096, p. 206.

Characters of the shell: Shell very much like that of L. viridis, but invariably
much smaller, thinner, and more delicate, with the hinge-teeth thin and weak.
The interdental projection of the left valve is quite rudimentary, and generally,
no indication of it is present. The outline of the shell is not so distinctly trape-
zoidal, but rather subovate, narrower in front, higher behind, and the upper
margin forms only a blunt angle with the posterior margin. The two valves are
not so distinctly compressed, as in L. viridis, and the swelling in the region of the
posterior ridge is generally more pronounced, so that the whole shell appears
relatively more inflated. Beak-sculpture similar to that of L. viridis, but the
bars are stronger and less in number (four); they do not extend so far upon the
disk, are more equal, with the posterior loop not so sharply angled (See Marshall,
1890, p. 176, fig. 2). Epidermis yellowish brown to olive greenish, with more or
less distinct green rays, which are most distinct and crowded upon the posterior
slope, so that the latter generally appears darker green. Nacre white and iridescent,
but often with lurid or dirty salmon tints.

122

MEMOIRS OF THE CARNEGIE MUSEUM.

L.

H.

D.

Size: I. Greencastle, Cat. No. 61.4222 (largest at hand).

. . 63 mm.

38 nim.

26 mm.

2. Yardley, Cat. No. 61.3689

..55 “

32 “

20 “

3. do. do.

..51 “

31 “

19 “

4. do. do.

..41 “

25 “

15 “

Soft parts (See Ortmami, 1912, p. 282). Glochidia (See Lea, Obs. XII, 1874,
PI. 21, fig. 4). Measurements: 0.36 X 0.30 mm.

This s]3ecies is also normally hermaphroditic, and specimens with male struc-
ture have never been found. Since the first publication of this fact I have ex-
amined several hundreds of additional specimens, but all had the female characters
in the gills.

Breeding season: Gravid females have been collected at the following dates:
Aug. 13, 1910; Aug. 20, 1909; Aug. 24, 1908; Sept. 5, 1909; Sept. 6, 1909; Sept.
12, 1912; Sept. 14, 1912; Sept. 16, 1912; then again: April 24, 1909; May 6,
1912; June 8, 1912. Thus the season begins about the middle of August. The
first date for the presence of glochidia is September 5. The glochidia are carried

• Lasmigona subviridis.

over the winter, and discharge was observed on the last date, June 8. The species
is bradytictic.

Remarks: Although this species morphologically is closely allied to L. viridis,
and is undoubtedly its eastern representative, and although the differences are
only in the degree of development of certain characters, it is not difficult to dis-
tinguish it at the first glance on account of the shape and size of shell. The range

ORTMANN: monograph of the naiades of PENNSYLVANIA.

123

of variation is not very great, and chiefly lies in the color of the epidermis. One
peculiar local phase, however, deserves special mention. In Conococheague Creek
at Greencastle, Franklin Co., this species is represented by rather large specimens,
which have an unusually heavy shell, with the nacre more frequently of a clear
salmon color, and the beaks in a beautiful state of preservation, even in large
individuals. This creek, at that locality, is so heavily charged with lime (fed by
big limestone springs), that a deposit of calcareous ooze is formed at the bottom,
and there is no doubt, that the peculiar local development is due to these local
conditions. Something similar, but not so strongly pronounced, has been observed
in Conedoguinet Creek at Carlisle; and in Virginia, in Reed Creek at Wytheville,
under the same conditions, a form has been observed, which corresponds entirely
to this form from Greencastle. i

Localities in Pennsylvania represented in the Carnegie Museuvn:

Delaware-drainage :

Delaware River, Yardley, Buck.s Co.; Shawnee, Monroe Co.

Schuylkill Canal, Manayunk, Philadelphia Co.

Susquehanna-drainage :

Conedoguinet Creek, Carlisle, Cumberland Co.

Dunning Creek (J. F. L. Raschen) and Raystown Branch of Juniata River (A. Koenig), Bedford, Bed-
for^l Co.

Chest Creek, Patton, Cambria Co.

Cush Cushion Creek, Green Township, Indiana Co. (D. A. Atkinson).

Chemung River, South Waverly, Bradford Co.

Potomac-drainage:

Conococheague Creek, Greencastle, Franklin Co.

Great Tonoloway Creek, Thompson Township, Fulton Co.

Localities in Pennsylvania represented in the Philadelphia Academy of Natural
Sciences:

“Valley Creek” southwest of Coatesville, Chester Co. (C. H. Conner).

Pennsylvania Canal, “ Chicques,”®° Pennsylvania (S. S. Haldeman).

Sinnemahoning Creek, Round Island, Clinton Co. (S. N. Rhoads).

Other localities represented in the Carnegie Museum:

Delaware-Raritan Canal, Princeton, Mercer Co., New Jersey.

South Branch Potomac River, Southbranch and Romney, Hampshire Co., West Virginia.

I found this species abundant in Little Antietam Creek, Waynesboro, Franklin Co., but the
specimens collected (about a dozen), on Aug. 10, 1910, were lost during the trip in an unaccountable way.

There is no creek of this name on the map, but “ Sucker Run ” is in the “ valley ” southwest
of Coatesville, tributary to West Branch Brandywine Creek. ^

No doubt “ Chickies,” near Columbia, Lancaster Co.

124

JMEMOIKS OF THE CARNEGIE MUSEUM.

Shenandoah River, Harpers Ferry, Jefferson Co., West Virginia.

South Fork Shenandoah River, Elkton, Rockingham Co., Virginia.

Rappahannock River, Remington, Fauquier Co., Virginia.

Rapidan River, Rapidan, Culpeper Co., Virginia.

North River, Buena Vista, Rockbridge Co., Virginia.

Kanawha-drainage :

Pool of Kanawha River, Glen Ferris, Fayette Co., West Virginia.

Greenbrier River, Ronceverte, Greenbrier Co., West Virginia.

New River, Hinton, Summers Co., West Virginia; Pearisburg, Giles Co., Virginia.

Reed Creek, Wytheville, Wythe Co., Virginia.

Distribution and Ecology in Pennsylvania (See fig. 13) : According to the
records at hand, this species is very erratic in its distribution. It is found in all
three drainages on the Atlantic side, but it is evident that it avoids the large
rivers and prefers smaller streams. Its absence . in the lists of Gabb (1861)^^ and
Schick (1895) is significant, although, as I have discovered, it is abundant in the
Schuylkill Canal at Manayunk. The specimens found by myself in larger rivers
generally were few, and often in small branches of the river.

But even in small streams, it is not everywhere present. But generally, when
found, it turned up abundantly. Like the preceding species (L. viridis), it is
averse to very strong current, and prefers more quiet parts, pools or eddies with
gravelly and sandy bottoms, and it also goes into canals, where it seems to flourish.
In the Susquehanna-drainage it goes far up into the headwaters, advancing west-
ward to Bedford, Cambria, and Indiana Cos.

Outside of Pennsylvania, it is found under similar conditions, and is also
more abundant in smaller streams. In the Kanawha-drainage it descends to the
big pool above the Kanawha falls, where it lives in an environment resembling a
lake.

General distribution: Type locality, Frankstown Branch Juniata River, Holli-
daysburg, Blair Co., Pennsylvania (Conrad).

Simpson (1900) gives: “streams draining into the Atlantic from New York
south to North Carolina,” and “Monroe Co., Michigan?”. Of course, this latter
locality, doubted already by Walker (1898), is certainly incorrect. We possess
the largest number of locality records, outside of Pennsylvania, from the state of
New York (Marshall, 1895), where it is found in the drainages of the Susquehanna
and Hudson Rivers, and it goes westwards through the Mohawk into the Erie
Canal, reaching the Genesee River (See Baker, 18986). As has been said above,
in this region it overlaps L. viridis, but particulars as to the mutual relations of
these two species are absent.

Gabb mentions it as a species, which might be found near Philadelphia. Conrad (1836) reports a
single individual from the Schuylkill at Philadelphia.

OKTMANN: monograph of the naiades of PENNSYLVANIA,

125

South of Pennsylvania records become scarce, and aside from those given
by myself we have only the following: Sideling Creek, Allegany Co., Maryland
(Pilsbry, 1894); Richmond, Henrico Co., Virginia (Lea’s U. hyalinus); and Neuse
River, Raleigh, Wake Co., North Carolina (Lea’s 17. pertenuis).

My discovery of the presence of this species in the upper Kanawha-drainage
in Virginia and West Virginia is quite astonishing and has been discussed elsewhere
(Ortmann, 1913a, p. 371, fig.). It should be added, that further investigations
have traced this species down to the Big Pool of the Kanawha (at Glen Ferris,
in Fayette Co., West Virginia).

Sub genus Lasmigona Rafinesque (1831).

Simpson, 1914, p. 488; Ortmann, 1914, p. 43.

Type Alasmidonta costata Rafinesque.

One species and one variety belong here.

Key to the Forms of Lasmigona.

ai. Shell compressed, not subcylindrical L. (L.) costata.

a2. Shell more swollen, and thus more nearly subcylindrical L. (L.) costata eriganensis.

Lasmigona (Lasmigona) costata (Rafinesque) (1820).

Symphynota costata (Rafinesque) Simpson, 1914, p, 488.

Plate IX, fig. 5.

Records from Pennsylvania:

Harn, 1891 (western Pennsylvania).

Stupakoff, 1894 (Allegheny Co.).

Marshall, 1895 (Allegheny River, Warren Co.).

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.; Beaver River, Wampum,
Lawrence Co.).

Ortmann, 19096, p. 196.

Characters of the shell: Shell rather large, moderately solid. Outline sub-
rhomboidal or subtrapezoidal, more or less elongated. Anterior end rounded,
posterior obliquely truncate. Lower margin gently convex, or almost straight.
Upper margin straight or slightly curved, somewhat ascending posteriorly, and
forming a rounded angle with the posterior margin. Beaks not prominent, placed
in front of the middle of the shell. Beak-sculpture consisting of about four or
five bars, which are rather heavy and swollen, the first simply curved, the others
with a tendency to fall into two loops, but this is distinct only in the second and
third bar, where the posterior loop is angled, and the sinus is present, but not

126

MEMOIRS OF THE CARNEGIE MUSEUM.

sharp. The later bars are only developed in the middle, and have only a slight
sinuation (see Marshall’s figure,- 1890, fig. 10). Shell rather flat, and more or less
compressed, with an indistinct, blunt posterior ridge. Greatest diameter in the
middle of the disk, or sbmewhat posterior to it, immediately in front of the posterior
ridge, and in this case the sides of the disk are flattened. Behind the posterior
ridge, the posterior slope is corrugated by heavy, blunt ridges or ribs, running
toward the posterior margin. In front of the posterior ridge, the disk is smooth,
without sculpture (sometimes there are short vertical grooves, such as are found
incidentally also in other Naiades).

Epidermis yellowish or brownish olive, greenish, brownish or blackish, with
more or less distinct, narrower or broader, dark green rays, which disappear in
old shells, which are more or less uniformly brownish or greenish black. Growth-
rests irregular, more or less distinct, often marked by darker bands.

Hinge partly obliterated. The pseudocardinals are moderately developed
(sometimes heavy in old shells), of the characteristic Anodontine type (more or
less lamellar and directed obliquely forwards), with the interdental projection
very well developed. Lateral teeth practically absent, or indicated only by mere
rudiments. Beak-cavity shallow. Dorsal muscle-scars in the beak-cavity. Nacre
white, often cream-color or pale salmon, chiefly so toward the beak-cavities.

No material sexual differences are seen in the shell. It has been asserted by
Scammon (1906) that the shell of the female is shorter and more inflated, while
Call (1900, p. 525) says that the females are a little more obese than the males.
According to my experience large specimens, which are rather swollen just in
front of the posterior ridge, are generally females (See Nos. 1 and 3 in table of
measurements) but there are females, which do not show this character (See No. 2),
and in medium-sized and small shells it is not at all developed (Compare Nos.
4-9) . As a rule, it is not easy to positively identify the sex by the shell alone, and
in younger shells this is impossible. That the females are shorter than the males
is not at all correct (See table of measurements).

L. H. D.

Size: 1. Enon Valley, Cat. No. 61.3134. 149 mm. 86 mm. 50 mm.

2. do. do. 137 “ 76 “ 37 “

3. Waynesburg, Cat. No. 61.4709 134 “ 77 “ 45 “

4. Harbison, Cat. No. 61.4710 126 “ 71 “ 35 “

5. Mount Morris, Cat. No. 61.4711 126 “ 67 “ 36 “

6. Waynesburg, Cat.' No. 61.4709 122 “ 72 “ 35 “

7. Harbor Bridge, Cat. No. 61.3709 122 “ 67 “ 32 “

8. Harbison, Cat. No. 61.4710 75 “ 41 “ 19 “

9. Wurtemberg, Cat. No. 61.4712 73 “ 41 “ 18 “

(swollen 9 , largest
at hand)

(flat 9 )

(swollen 9 )
(swollen cf)

(9)

id')

(flat 9 , gravid!)
(c^)

(9)

ORTMANN; monograph of the naiades of PENNSYLVANIA.

127

Soft parts (See Ortmann, 1912, p. 283). Glochidia (See Lea, Obs. VI, 1858,
PL 5, fig. 26; Lefevre & Curtis^ 1910, p. 97, fig. B, and 1912, p. 146, fig. B; Surber,
1912, PI. 1, fig. 7). Surber gives the following dimensions: 0.385 X 0.390; Lefevre
& Curtis; 0.35 X 0.39; while I (1912) give: 0.34 X 0.37 mm.

Breeding season: 1 have the following records for gravid females: August 3,
1909; Aug. 8, 1914; Aug. 9, 1907; Aug. 23, 1916; Aug. 29, 1910; Aug. 31, 1906;
Sept. 2, 1907; Sept. 2, 1908; Sept. 4, 1908; Sept. 5, 1908; Sept. 5, 1913; Sept. 7,
1908; Sept. 7, 1913; Sept. 10, 1906; Sept. 11, 1913; Sept. 12, 1913; Sept. 13,
1910; Sept. 13, 1915; Sept. 15, 1913; Sept. 17, 1912; Sept. 17, 1917; Sept. 18,
1917; Sept. 21, 1907; Sept. 21, 1908; Sept. 27, 1909; Sept. 28, 1911; Oct. 4, 1910;
Oct. 14, 1907; Oct. 15, 1907; Oct. 19, 1908; Oct. 23, 1907. Then again in spring:
April 22, 1908; April 24, 1908; May 11, 1907; May 11, 1911; May 13, 1910;
May 17, 1910; May 26, 1908.

Thus this species is hradytictic. The breeding season begins early in August,
and by the end of this month (earliest date Aug. 31), and in September, glochidia
begin to be present. They are carried over the winter, and are found in the mar-
supium in April and May, and are discharged during May (observed on Alay 11
and 13). In June and July, there is an “interim,” and although many specimens
were collected, there never was in these months a gravid female among them.

Remarks: This species is easily recognized by the general shape, sculpture of
the posterior slope, and conformation of the hinge. From L. viridis, which it
resembles somewhat in shape, it differs also in beak-sculpture. Variations are chiefly
due to differences in the relation of length to height, and the development of the
sculpture of the posterior slope, which may be coarser or finer, and may be more or
less distinct; but traces of this sculpture are always present. I have not been
able to discover any marked tendency in the direction of the development of local
races, unless it be in size, the forms of certain creeks always remaining compara-
tively small. This is especially true of some little streams in the mountains (upper
Loyalhanna River in Westmoreland Co., Quemahoning Creek in Somerset Co.,
Little Mahoning Creek in Indiana Co.). Other small creeks, chiefly in the north-
western part of the state, favor the development of almost gigantic forms, for
instance. Little Beaver Creek, the upper Shenango River, and others.®^

1 found outside of our state in the upper Tygart River at Elkins, Randolph Co., West Virginia, a
rather interesting local form, with a very pronounced tendency toward the obliteration of the sculpture
of the posterior slope, which in these specimens in most cases is perfectly smooth, although individuals
occur, which show rudiments of the ribs.

128

MEMOIRS OF THE CARNEGIE MUSEUM.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River and its smaller tributaries:

Ohio River, Industry, Beaver Co.; Dead Man’s Island, Allegheny Co. (R. Foerster); Neville Island,
Allegheny Co.

Buffalo Creek, Acheson, Washington Co.

Little Beaver Creek, Cannelton (Miss Vera White, H. H. Smith), Darlington and New Galilee, Beaver
Co.; Enon Valley, Lawrence Co.

Raccoon Creek, New Sheffield, Beaver Co.; Bavington, Washington Co.

Chartiers Creek, Carnegie, Allegheny Co. (D. A. Atkinson).*®

Beaver-drainage:

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Ellwood City, Lawrence Co. (G. 11. Clapp & H. 11. Smith); Harmony, Butler Co.
Slipperyrock Creek, Wurtemberg and Rose Point, Lawrence Co.

Wolf Creek, Grove City, Mercer Co.

Little Connoquenessing Creek, Harmony, Butler Co.

Vlahoning River, Mahoningtown, Coverts, Edinburg and Hillsville, Lawrence Co.

Neshannock Creek, Volant, Lawrence Co.; Leesburg, Mercer Co.

Otter Creek, Mercer, Mercer Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsville, Clarksville, Shenango, and
Jamestown, Vlercer Co.; Linesville, Crawford Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

Little Shenango River, Greenville, Mercer Co.

Allegheny-drainage:

Allegheny River, Godfrey, Kelly, Mosgrove, and Templeton, Armstrong Co.; Walnut Bend, Venango
Co.; Tionesta and Hickory, Forest Co.; Warren, Warren Co.; Eldred, VIcKean Co.; Larabee,
McKean Co. (Dennis Dally).

Buffalo Creek, Harbison, Butler Co.

Loyalhanna River, Idlepark and Ligonier, Westmoreland Co.

Two Lick Creek, Homer, Indiana Co.

Quemahoning Creek, Stantons Mill, Somerset Co.

Crooked Creek, Rosston and South Bend, Armstrong Co.; Creekside, Indiana Co.

Cowanshannock Creek, Rural Valley, Armstrong Co. (N. M. Grier).

Little Mahoning Creek, Goodville, Indiana Co.

Sandy Creek, Sandy Lake, IMercer Co.

French Creek, Utica, Venango Co.; Cochranton and Meadville, Crawford Co.

Sugar Creek, Cooperstown, Venango Co. (E. P. Spencer).

Conneauttee Creek, Edinboro, Erie Co.

Leboeuf Creek, Waterford, Erie Co.

Brokenstraw Creek, Garland, Warren Co.

Connewango Creek, Russell, Warren Co.

Potato Creek, Smethport, McKean Co. (P. E. Nordgren).

M onongahela-drainage:

Monongahela RWer, Elizabeth, Allegheny Co. (D. A. Atkinson); Charleroi, Washington Co. (G. A.
Ehrmann); Westmoreland Co. (G. A. Ehrmann).

*® Dead shell seen in Little Chartiers Creek, Morganza, Washington Co.

ORTMANN: monograph of the naiades op PENNSYLVANIA.

129

Ten Mile Creek, Amity, Washington Co.

South Fork Ten Mile Creek, Waynesburg, Greene Co.

Dunkard Creek, Wiley and Mount Morris, Greene Co.

Cheat River, Cheat Haven, Fayette Co.

Localities in Pennsylvania represented in the Philadelphia Acade7ny of Natural
Sciences:

Ohio River, Beaver, Beaver Co., and Coraopolis, Allegheny Co. (S. N. Rhoads).

Other localities represented in the Carnegie Museum:

Atlantic- and Lake-drainage:

Mohawk River, New York (Smith collection).

Sandusky River, Upper Sandusky, Wyandot Co., Ohio (C. Goodrich).

Miami and Erie Canal, Waterville, Lucas Co., Ohio (C. Goodrich).

Maumee River, Waterville, Lucas Co., and Defiance, Defiance Co., Ohio (C. Goodrich).

St. Marys River, Rockford, Monroe Co., Ohio (C. Goodrich).

Raisin River, Grape P. 0., Monroe Co., Michigan (C. Goodrich).

Kaministiquia River, Stanley, western Ontario, Canada (0. E. Jennings).®^

Ohio-dramage:

Ohio River, Portsmouth, Scioto Co., Ohio.

Wolfe Creek, Washington Co., Ohio (W. F. Graham).

Scioto River, Kenton, Hardin Co., Ohio (C. Goodrich).

Rock River, Illinois (Smith collection).

Cheat River, Jaco and Mount Chateau, Monongalia Co., West Virginia.

West Fork River, Lynch Mines, Harrison Co.; Lightburn and Weston, Lewis Co., West Virginia.
Tygart River, Elkins, Randolph Co., West Virginia.

Little Kanawha River, Grantsville, Calhoun Co. ( W. F. Graham) ; Burnsville, Braxton Co., West Virginia.
North Fork Hughes River, Cornwallis, Ritchie Co., West Virginia.

Elk River, Sutton and Gassaway, Braxton Co.; Shelton, Clay 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).

Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).

Shoals Creek, Lauderdale Co., Alabama (H. H. Smith).

Elk River, Fayetteville, Lincoln Co., and Estill Springs, Franklin Co., Tennessee (H. H. Smith).

Flint River and Hurricane Creek, Gurley, Madison Co., Alabama (H. E. AVheeler).

Paint Rock River, Trenton and Princeton, Jackson Co., Alabama (H. H. Smith).

South Chickamauga Creek, Ringgold, Catoosa Co., Georgia.

Tennessee River, Knox Co., Tennessee (Smith collection).

Boyd Creek, Boyd Creek, Sevier Co., Tennessee.

Holston River, Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill, Noeton, and Holston Station,
Grainger Co.; Austin Mill and Church Hill, Hawkins Co., Tennessee.

About 15 miles west of Fort Williams on the Kaministiquia River, which flows to Lake Superior,
but it might have a connection with the Hudson Bay-drainage. Special attention should be called to
this new locality !

130

MEMOIRS OF THE CARNEGIE MUSEUM.

South Fork Holston River, Pactolus, Bluff City, and Emmett, Sullivan Co., Tennessee.

Middle Fork Holston River, Chilhowie, Smyth Co., Virginia.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia; Mendota,

Washington Co., Virginia; Saltville, Smyth Co., Virginia.

Clinch River, Edgemoor and Offutt, Anderson 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; Raven, Richland, and Cedar Bluff, Tazewell

Co., Virginia.

Powell River, Combs, Claiborne Co., Tennessee; Dryden, Lee Co., Virginia.

Distribution and Ecology in Pennsylvania (See fig. 12) : This species is widely
distributed in the headwaters of the Ohio in western Pennsylvania, and is found
in large rivers as well as in very small creeks, although it is distinctly more abundant
in the latter. There is hardly any small stream from which it is entirely absent,
but in the large rivers, although present, it is decidedly rare.

The wide and general distribution of this species indicates that it is not very
particular as to station. It is found under a great variety of conditions, in riffles,
with a strong and variable current, in steady, stronger or weaker currents, and
even in rather quiet water and deep pools. It is found on every variety of bottom,
but prefers coarser or finer gravel, and more rarely is seen in sand and mud. It is
one of the few species, which advances far eastwards in the headwaters of the
Conemaugh and Allegheny drainages, but although it closely approaches the divide,
it has never been found in Pennsylvania to the east of it in the Atlantic-drainage.

It has not been found in the only tributary of Lake Erie in our state, which
has shells (Conneaut Creek).

General distribution: Type locality, Kentucky River (Rafinesque) .

This species has an enormous range. It is found practically all over the
Mississippi and Ohio-drainages, from western Pennsylvania throughout Ohio,
Indiana, Illinois, West Virginia, Kentucky, and Tennessee to northern Alabama.
In the upper Tennessee-drainage it goes by the Powell, Clinch, and Holston Rivers
into Virginia. In a southwesterly direction it extends to eastern Kansas, through
Arkansas to Oklahoma and northern Louisiana.

Simpson cites it from Columbus, Lowndes Co., Mississippi, in the Tombigbee-
drainage, but this is the only place from which it has been recorded in the Alabama
system.

Westward it does not seem to go beyond Iowa (abundant in the Wapsipinicon,
Volga, and Turkey Rivers, in northeastern Iowa, according to Geiser, 1910), but
in a northerly direction it has crossed over into the Hudson Bay-drainage in Mani-
toba, and possibly our locality in a northern tributary of Lake Superior (Kam-
inistiquia River), is connected with this region,

ORTMANN: monograph of the naiades of PENNSYLVANIA.

131

In the drainage of the Great Lakes, from Wisconsin through Michigan to New
York and eastern Canada it is widely distributed. However this latter part of
its range should be investigated more closely. As we shall presently see, in Lake
Erie a peculiar local variety has been developed. Nevertheless in the lake-drainage
in western and central New York the normal type is present. Here it advances
along the Erie Canal to the Mohawk and even to the Hudson River. It follows
the lower St. Lawrence down to the neighborhood of Ottawa and Montreal (and
also occurs in the Lake Champlain region), but it is not known whether the form
peculiar to Lake Erie or the normal type prevails here.

It should be further noted that the rule that this species is rare in large rivers
holds good also in the Ohio below Pittsburgh. Between the Pennsylvania state-
line and Cincinnati, I found only a single individual (in a clam-digger’s pile at
Portsmouth, Scioto Co., Ohio). This is in strong contrast to the abundance of
this species in some of the smaller creeks in West Virginia and Kentucky.

Lasmigona (Lasmigona) costata eriganensis Grier (1918).

A form, not distinguished hitherto, except by Sterki (1907a, p. 393) who called attention to its
peculiarities, and by Grier (1918, p. 10) who elevated it to the rank of a variety.

Plate IX, fig. 6.

Characters of the variety: Shell smaller, slightly more elongate, more swollen,
and thus approaching in a degree the subcylindrical shape. Ridges of posterior
slope generally narrower. Color lighter, more yellowish-brown, sometimes rusty
brown, with the rays indistinct, and the growth-rests more distinct and more

regular.

L.

H.

D.

Size: 1.

Presque Isle Bay, Cat. No. 61.4223 .

. 90 mm.

44 mm.

31 mm.

Type Set

2.

do.

do.

.88 “

45 “

26 “

U li

3.

do.

do.

.85 “

46 “

29 “

{( (C

4.

do.

do. .

.76 “

39 “

28 “

(C cc

5.

do.

do.

.73 “

38 “

25 “

C( u

Compared with the measurements of typical costata (p. 126), the differences
in shape become evident, especially the greater diameter.

Soft parts: Living specimens have been found only on two occasions: May 22,
1909; and July 8, 1910. Although there were females among them, none were
gravid. The soft parts of none were preserved, but as far as I am able to recol-
lect, they were not different from the normal form.

Remarks: Although I have only fifteen specimens from Pennsylvania, and
five from Michigan (to which might be added about half a dozen individuals used

132

MEMOIRS OF THE CARNEGIE MUSEUM.

in exchange) I feel quite sure, that Grier is right in thinking that this shell repre-
sents a very peculiar phase of L. costata. The whole outer aspect of this shell is
quite unlike that of the normal form, but, of course, the sculpture of the posterior
slope and the hinge do not leave any doubt as to its systematic relationship. As
the above measurements show, there is some variation in shape, and furthermore
the color of the epidermis may be lighter or darker, sometimes inclining more to
brown, sometimes to rusty. Greenish tints are rarely seen.

Localities represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania.

Lake Erie, La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).

Distribution and Ecology (See fig. 12). Type locality: Lake Erie; Presque
Isle Bay, Erie, Erie Co., Pennsylvania. Type set: Carnegie Museum Cat. No.
61.4223 (not 61.4720, as Grier states!).

I know this form only from the Lake Erie shores in Pennsylvania and Michi-
gan, but according to Sterki it also occurs on the Ohio shores, and very likely will
be found elsewhere in Lake Erie. Walker’s (1913, ]). 22) S. costata from Lake Erie
is undoubtedly this form. The specimens collected by myself are all from the
north shore of the bay “Big Bend,” where I found them alive in one to three feet
of water, on sand and fine gravel, without vegetation, and at places, which are
exposed at times to a considerable surf. This form is not abundant there. Some
specimens were found among a scanty growth of rushes {J uncus americanus).

Note: Since it is possible that L. costata of western New York may have come
by way of Lake Erie, it is interesting to observe that the typical L. costata is found
in this region, as indicated by De Kay’s figure (1843, PI. 14, fig. 226) from Oswego
River (Lake Ontario-drainage) . This possibly is a case parallel to others men-
tioned above, where the lake-form, when migrating into the tributaries, assumes
again the shape of the normal creek-form. The matter, however, is not perfectly
clear, for there is the possibility that western and central New York was reached
by way of the upper Allegheny.

Subgenus Pterosyna Rafinesque (1831).*^

Pterosygna Simpson, 1914, p. 490; Ortmann, 1914, p. 43.

Type Unio complanatus Barnes.

Only one species is known to belong to this subgenus.

Pterosyna, and not Pterosygna^ as Simpson writes.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

133

Lasmigona (Pterosyna) complanata (Barnes) (1823).

Symphynota complanata (Barnes) Simpson, 1914, p. 490.

Plate IX, fig. 7.

Records from Pennsylvania :

Ortmann, 19096, p. 196.

Characters of the shell: Shell large; rather thin when young, moderately solid
when old. Outline broadly ovate, subrhomboidal, not elongate, only a little longer
than high. Anterior end rounded, posterior obliquely truncate. Lower margin
convex, less so posteriorly. Upper margin nearly straight, strongly ascending
posteriorly, forming a distinct angle with the posterior margin, and projecting so
as to form a vexy distinct wing, the upper margin of which is symphynote. The
wing is best developed in young specimens. Beaks not. prominent, placed in front
of the middle of the shell. Beak-sculptiHe consisting of four to five rather distinct
and sharp bars, the first one or two simply curved, the others strongly double-
looped, with the anterior loop round and wide, the posterior narrower and angled,
and with a sharp, re-entering angle between them. Shell flat and greatly com-
pressed, with an indistinct, blunt and broad posterior ridge. The posterior slope
is alate and strongly compressed, and in many cases is ornamented with irregular
ridges running toward the posterior margin, which, however, may be entirely
absent. Disk in front of posterior ridge smooth, without sculpture.

Epidermis yellowish to greenish brown; when young with indistinct traces of
green rays, which become entirely obsolete in older shells, of which the epidermis
is lighter or darker brown, often blackish. Sometimes there is some dark green in
the epidermis. Growth-rests more or less distinct, often marked by darker color.

Hinge-teeth partly obliterated. Pseudocardinals present and rather heavy,
and quite variable in shape. Interdental projection variable, sometimes rather
distinct; in other cases weak, and often entirely absent. Lateral teeth practically
absent, indicated only by vestigial blunt ridges. The conformation of the hinge
in this species is extremely and remarkably variable. Beak-cavity shallow.
Dorsal muscle-scars in the beak-cavity. Nacre white, sometimes lightly tinted
with cream-color toward the beak-cavity.

No differences in the shape of the shell in the two sexes. Baker (1898a,
p. 60) and Scammon (1906, p. 332) say that the male is more compressed than the
female. This is not at all correct, as is easily seen by comparing nos. 2, 3, and 4,
and nos. 5 and 6 in our table of measurements.

134 ^ MEMOIRS OF THE CARNEGIE MUSEUM.

L. H. D.

Size: I. Conneaut Outlet, Cat. No. 61.3315 (c?) 124 mm. 93 mm. 40 mm.

2. Waterford, Cat. No. 61.4238 (d") 118 “ 86 “ 40 “

3. do. “ “ do. (9)....' 118 “ 84 “ 39 “

4. Conneaut Outlet, Cat. No. 61.3315 (9) 117 “ 88 “ 35 “

5. do. " “ do. (9) 93 “ 71 “ 23 “

6. Waterford, Cat. No. 64.4237 (d") 90 “ 65 “ 25 “

7. do. Cat. No. 61.4238 (c?) 78 “ 60 “ 21 “

Outside of Pennsylvania this species grows much larger.

Soft parts: Figured by Lefevre & Curtis, 1910, PI. 1, fig. 6, and 1912, PI. 6,
fig. 3; described by Ortmann, 1912, p. 282. Glochidia (See Lea, Obs. VI, 1858,
PI. 5, fig. 29; Lefevre & Curtis, 1910, p. 97, fig. A, and 1912, p. 146, fig. A; Ort-
mann, 19115, PL 89, fig. 11; Surber, 1912, PI. 1, fig. 6. Lefevre & Curtis give the
dimensions: 0.28 X 0.30 mm.; Surber: 0.310 X 0.320; while I gave: 0.34 X
0.34 mm.

Breeding season: I found gravid females on September 14, 1909; September
22, 1910; and May 14, 1908. Glochidia were observed on the first and last dates.
Although these records are very meager, yet they distinctly indicate that we have
to deal with a bradytictic form, which carries the- glochidia over winter. According
to Surber (1912, p. 7), this species is gravid in October, November, and May.

Remarks: An extremely characteristic species, which in its external shape
greatly resembles Proptera alata (Say), but may be distinguished at once by the
color of the nacre (white, not purple), by the hinge, and some minor characters.
The similarity of the outer shape of these two species, belonging to different sub-
families, is very remarkable, and offers a very good example of parallel, but inde-
pendent, development of the same external characters (“convergency ”).

Aside from slight differences in outline and the variability of the hinge, this
species varies chiefly in the development of the sculpture of the posterior slope.
In Pennsylvania both conditions, specimens with well-developed sculpture, and
with practically smooth shells, are found side by side, accompanied by all tran-
sitional stages. In size the Pennsylvanian shells remain far below the dimensions
recorded for specimens from farther west (Scammon gives: L. 188; H. 110: D.
52 mm.).

Localities in Pennsylvania represented in the Carnegie Museum:

Conneaut Outlet, Conneautlake, Crawford Co.

Lcboeuf Creek, Waterford, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

St. Josephs River, Fort Wayne, Allen Co., Indiana (C. Goodrich).

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135

Ohio-drainage:

Ohio River, Wheeling, Ohio Co., West Virginia (W. F. Graham); Portland, Meigs Co., Ohio.®'’

Wolfe Creek, Wolfe Creek P. 0., Washington Co., Ohio (W. F. Graham).

West Fork White River, Riverside, Green Co., Indiana (J. D. Haseman).

Wabash River, New Harmony, Posey Co., Indiana (A. A. Hinkley).

Aliddle Island Creek, Union Alills, Pleasants Co., West Virginia.

Pocatalico River, Rajanond City, Putnam Co., West Virginia.

West of Mississippi:

Roche-Perche Creek, Columbia, Boone Co., Alissouri (D. K. Greger).

Kansas River, Lawrence, Douglas Co., Kansas (R. L. Moodie).

Solomon River, Salina, Saline Co., Kansas (R. L. jMoodie).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).

Chikaskia River, Tonkawa, Kay Co., Oklahoma (F. B. Isely).

Alabama-drainage:

Big Prairie Creek, Hale Co., Alabama (Dr. Showalter, Hartman collection).

Cahaba River, Gurnee, Shelby Co., Alabama (H. H. Smith).

Distribution and Ecology (See fig. 12): Type-locality, Fox River, Wisconsin
(Barnes) .

The distribution of this species in Pennsylvania is very remarkable. It is
found only in two places; in both in the outlet of a glacial lake. At Conneaut
Outlet I found this species in rather strongly flowing water, with a soft bottom of
muddy sand; in Leboeuf Creek in a soft shell-marl. At both places the size of
the stream is about the same, rather small, with the supply of water uniform,
regulated by the lake above.

While at both localities the species is quite frequent, no trace of it has ever
been found in any other stream in this state. Down the Ohio, it turns up for the
first time at Wheeling, and further down at St. Marys, Parkersburg, and Portland.
But at St. Marys it is not in the Ohio proper, but in the slackwater of Middle
Island Creek, which goes up about five miles (to Union Mills). In the Kanawha-
drainage it goes up to Pocatalico River, in the lower part of which it is not rare,
being chiefly found in quiet and muddy pools.

L. complanata has been reported to prefer deep and quiet water with muddy
bottoms (Baker, 1898a, Call, 1900, Scammon, 1906) although Scammon mentions
that it is occasionally found in riffles and in gravel. I have found it under such
conditions in the smaller branch of the Ohio at Portland, but only a single indi-
vidual. The lack in Pennsylvania of the environment it prefers accounts for its
absence in the upper Ohio-drainage in general. So much more astonishing is its
presence in the outlets of two glacial lakes, and I am not prepared to say, which

I liave also seen dead shells at Parkersburg, Wood Co., West Virginia, in the small branch of the
Ohio at Neal Island.

136

MEMOIRS OF THE CARNEGIE MUSEUM,

are the environmental features, which have favored its establishment there. Fur-
thermore the question remains to be answered how it succeeded in reaching these
isolated stations.

With regard to this phenomenon, attention may be called to the fact that this
species, although distributed over the whole state of Ohio (Sterki, 1907a), is especi-
ally found in its northeastern part, not far from the Pennsylvanian localities.
Dean reports it from Mahoning River; Silver Creek, Portage Co!.; and from
Cuyahoga River (lake-drainage). It seems that there might be here a connection
with the Pennsylvanian localities, and then this species very likely would belong
in the same category as Lasrnigona viridis (see above).

This sjiecies has a very wide distribution, and is found according to Simpson
(1900) in the “upper Mississippi-drainage as far south as Arkansas in the west;
Ohio River-system; upper St. Lawrence and its tributaries, north into Mackenzie
River.” A number of localities are known in the Lake Winnipeg and Nelson
drainages; in Lake of the Woods belonging to the upper Lakes drainage. It
occurs also in Michigan (Walker, 1898) in the lake-drainage, and in Rouge River
at Detroit belonging to the Lake Erie-drainage. It is found in the Erie-drainage
in Indiana and Ohio (Goodrich, 1914, Dean), but it has never been reported from
Lake Erie proper (Walker, 1913, p. 22), A record from Buffalo cannot be credited
on account of the general inaccuracy of this list, and Calks “western New York”
remains unconfirmed (Marshall, 1895). In addition, it goes southward into the
Alabama-drainage, according to Call (1885) and Lewis (1877) (See also specimens
in Carnegie Museum). Strangely enough localities from South of the Ohio in
Kentucky and Tennessee are not known, except the record of Wilson & Clark
(1914) from the Cumberland in Tennessee. Apparently, the Alabama localities
are connected with the rest of the range not in a northerly, but in a westerly direc-
tion. The species is found in northern Louisiana (Marshall, 1895; Vaughan, 1893).
It extends westward to Iowa, Kansas, Arkansas, and Oklahoma.

Note: SiMPSONicoNCHA AMBiGUA (Say) (1825).

Hemilastena ambigua (Say) Simpson, 1914, p, 325.®^

This species has never been found in Pennsylvania. According to Simpson
it is distributed in the “Ohio River system; north to Michigan; west to Iowa;
south to Arkansas; east to Tennessee.” According to Dean (1890), it is rare in
the Alahoning River, Ohio, but I have hunted for it in vain in the Pennsylvanian
part of this river.

No trace of it in the Pennsylvanian part of this river.

As to the generic names Simpsonaias and Simpsoniconcha, see Frierson, I9I4a, p. 7, and 1914&,

p. 40.

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137

On May 23, 1912, I found a single specimen of this species in the headwaters
of the Monongahela in West Fork River, Lightburn, Lewis Co., West Virginia.
I found it buried deeply in sand and gravel held together by the rhizomes and roots
of Dianther a.

Being thus present in two rivers, which run into our state, it might have once
existed here, or else it accidentally may have escaped detection.

The systematic position of this genus is as yet quite uncertain. The account
of the anatomical structure given by Simpson is quite unsatisfactory, in fact
partly unintelligible. Besides the nomenclature of the genus requires final adjust-
ment.

Genus Anodonta Lamarck (1799).

Ortmann, 1912, p. 286; Simpson, 1914, p. 358.

Type Mytilus cygneus Linnaeus.

A large number of species of this genus are known, many of which may be
only special phases of others. In Pennsylvania, four species and one or two
varieties may be distinguished, but I am not sure whether all of them are ‘‘good
species.”

Key to the Species and Varieties of Anodonta in Pennsylvania.
a I. Umbonal region convex, or more or less inflated and elevated above the hinge-line. Beak-sculpture
consisting of several rather strong, double-looped bars, which generally have a sharp re-entering
or angular sinus. Gonochoristic.

bi. Beak-sculpture rather heavy, forming distinct nodules on the posterior loop.

Cl. Shell rather compressed, not swollen.

di. Shell not very thin, ovate, somewhat elongate.. . A. grandis.

do. Shell rather thin, large, higher and shorter A. grandis gigantea.^^

C2. Shell more swollen and inflated, chiefly in the umbonal region A. grandis footiana.

62- Beak-sculpture less heavy, generally not nodulous, and bars of uniform thickness.

Cl. Shell rather thin or moderately thick, not thickened along the lower anterior margin.

A. cataracta.

C2. Shell somewhat thicker, and especially so along the lower anterior margin A. implicata.

a2. Umbonal region remarkably depressed and flattened, and not elevated above the hinge-line. Beak-
sculpture consisting of several delicate, irregular bars, with a slight tendency to fall into two loops,
the sinus being a mere notch. Hermaphroditic A. ohiensis.

I shall not treat this as a “ variety.” It has been introduced here only to bring out its characters.

138

MEMOIRS OF THE CARNEGIE MUSEUM.

Anodonta grandis Say (1829).

Anodonta grandis Say; Simpson, 1914, p. 418.

Plate X, figs. 1, 2.

Records from Pennsylvania:

Ortiuann, 19096, p. 195. It is very remarkable that this common species has never before been reported
from western Pennsylvania.

Characters of the shell: Shell large, sometimes very large, but generally rather
light and thin, often very thin. Outline subovate or subelliptical, more or less
elongate, rounded anteriorly, more or less narrowed behind, often somewhat
pointed. Lower margin more or less convex, upper margin straight, often forming
an angle with the obliquely descending posterior margin, which in many cases
forms a moderately developed wing, but in other cases may be altogether absent.
Young shells are often symphynote at the upper posterior angle. Beaks slightly
swollen and moderately convex, somewhat elevated above the hinge-line, placed
more or less anterior to the middle of the shell. Beak-sculpture (Marshall, 1890,
figs. 15, 18, as of A. lewisi) normally distinct and rather heavy, consisting of four
to five (rarely six) bars, of which the first and second are concentric, while the
following are distinctly double-looped, with a sharp re-entering sinus between the
loops. The anterior loop is broadly rounded, while the posterior is angular,
narrower, and characteristically elevated, so as to form a distinct tubercle. How-
ever, there is great variability in the beak-sculpture, chiefly with regard to the
development of the sinus and the tubercle. The former may be shorter or longer,
often so long as to reach the preceding bar; often the bar is weak at the sinus, so
that a notch is formed, the anterior and posterior loop being higher than this part.
The tubercles may be more or less distinct, and sometimes also the posterior end
of the anterior loop (close to the sinus) is tubercular, so that a double row of tub-
ercles appears to be present, radiating from the beaks. In partly eroded beaks,
often only the tubercles are preserved.

Shell gently swollen in the middle and toward the beaks, but swelling very
variable, sometimes rather flattened on the disk. Posterior ridge indistinct,
greatest diameter of the shell about the middle. Posterior slope slightly com-
liressed, often elevated and winged at the upper posterior angle.

Epidermis yellowish, greenish, to brownish and blackish. Rays indistinct as
a rule; in large specimens (and often also in younger ones), there is no trace of
rays on the disk. In other cases there are traces of gveen rays, but they are never
very distinct, and, when present, generally obliterated by the concentric color-

Simpson (1900, p. 644) gives, with ??, southeastern Pennsylvania, but this is surely incorrect.

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139

pattern. The latter consists of concentric bands of lighter and darker color,
largely corresponding to growth-periods. Upon the lighter bands the rays are
sometimes visible. However, very often there are one to three more distinct,
dark green to black rays upon the posterior slope, which are best visible in trans-
mitted light. In many specimens, chiefly in the common creek-form, the epidermis
is uniformly blackish.

Hinge edentulous, forming a practically straight line, with only the faintest
trace of a light undulation under the beaks. Beak-cavity shallow or moderate.
Dorsal muscle-scars faint, situated in the beak-cavity. Nacre white, as a rule,
silvery, bluish white, or cream-color, rarely pinkish, generally highly iridescent,
with blue, purple, and greenish reflections. In some cases it is more distinctly
reddish (salmon to rusty), but this is pathological (see below under salmonia).

There is a difference in the shape of the shell of the two sexes, but this is very
slight, and not always present. In the females, and chiefly in the larger ones, the
shell is more swollen in the posterior middle portion of the disk, just in front of
the indistinct posterior ridge, in the region where the marsupium is located. This
swelling sometimes renders the lower margin more curved and slightly prominent
behind the middle. In extreme cases the shell is even slightly depressed in front
of the swelling. In young and half-grown shells this shape is not noticeable, and
even older shells do not always show it. Where it is distinctly developed, it can
be safely taken for an indication of the female sex, but not all females have this
character. It also should be said, that in the typical (creek-) form, this sexual
difference is much less evident, than in the form of quiet water (gigantea).

L. H. D.

Size: 1. Linesville, Cat. No. 61.845 (largest on hand) .... 128 mm. 70 mm. 44 mm.

2. Edinboro, Cat. No. 61.3648 (9 gravid) 126 “ 62 “ 44 “

3. Waynesburg, Cat. No. 61.4729 (9) 123 “ 68 “ 40 “

4. Cochranton, Cat. No. 61.3661 (9 gravid) 118 “ 68 “ 39 “

5. Jamestown, Cat. No. 61.3659 (cf) 105 “ 61 “ 41 “

6. Waynesburg, Cat. No. 61.4729 (d") 91 “ 52 “ 33 “

Soft parts (See Ortmann, 1912, p. 292). Glochidia figured by Lea, Obs. VI,
1858, PL 5, figs. 32-34 and Surber, 1912, PL 3, fig. 45. Surber’s measurements
are: 0.41 X 0.42, while I gave: 0.36 X 0.37 mm.

Breeding season: Gravid females have been frequently found from August 6
to November 12, and again on May 22. This species seems to discharge very
early in spring (April). May 22 seems to be an exceptionally late date.

Remarks: This is a very variable species, for which a great number of specific
names has been introduced, chiefly by Lea. The typical form (of which ovata

140

MEMOIRS OF THE CARNEGIE MUSEUM.

Lea, lewisi Lea, salmonia Lea, harpethensis Lea simply are synonyms) is a shell
preferably inhabiting small creeks. It is chiefly characterized by its rather elon-
gated, subovate shape, with rather flat valves, by the comparative solidity of the
shell, its dark (greenish to blackish) color, and the nodulous character of the beak-
sculpture. The posterior end of the shell generally is narrowed, and the posterior
point is moderately elevated above the base line; and there is hardly a wing (how-
ever, this is sometimes indicated in the young). Yet all these characters are
variable, and certain extreme forms are found under peculiar conditions of environ-
ment, so that they are to be regarded as ecological variations, which turn up,
within the range of the species, wherever the proper conditions are found. A
number of them have received names; but these names should not be used as
varietal names, since the forms designated by them have no defined geographical
areas. They certainly are not ‘‘subspecies” in the strict taxonomic sense. I shall
mention here some of these “forms,” as far as they are found in Pennsjdvania,
with the express understanding, that I regard them only as individual variations,
or at the utmost as special reactions to peculiar environmental conditions, which
are not regionally restricted.

Anodonta grandis forma salmonia (Lea) (Obs. II, 1838, PL 14, fig. 41).

This is an unqualifiedly typical grandis, with the nacre peculiarly discolored
(reddish, from pale salmon to deep rusty red), and at the same time the nacre
liecomes rough by the formation of small pustules and granulations. That this
is due to an infection by a parasite (a distomid) is now generally conceded (see
Wilson & Clark, 1912a, p. 47), and consequently the name salmonia has no tax-
onomic standing, and might lietter be entirely dropped.

In Pennsylvania, such diseased forms are rather frequent in the northwestern
corner of the state (headwaters of Beaver system, and small tributaries of the
upper Allegheny), but are absent in the southwestern section. Sometimes traces
of this disease are found in pond- and lake-forms. At certain localities, practically
all individuals are more or less infected with this parasite.

Anodonta grandis forma gigantea (Lea) (1834) (Simpson, 1914, p. 420).

This is the form found in small, muddy pools, where it lives deeply buried in
black muck. It is distinguished from the creek-form by its larger size, thinner,
and higher and less elongated shell, and generally by more vivid color (yellowish
and greenish concentric bands, and more distinct traces of rays). However, this
form also is very variable, even in the same pond, and it passes insensibly into the
creek-form. In ponds more elongated specimens are often found, and in creeks
specimens from quiet and muddy pools distinctly incline toward the pond-form.
Specimens from ponds often also possess a rather thick shell.

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141

This form is rather scarce in Pennsylvania. My supply of it comes chiefly
from a pond (old branch of Allegheny River) at Harmarville, Allegheny Co. But
other specimens are at hand from an artificial ice pond at Edgeworth, Allegheny
Co., and from a pond at Idlepark, Westmoreland Co., formed by the cutting off of
a part of Loyalhanna River by the railroad. Probably this form will turn up in
other ponds of the western part of the state.

L. H. D.

Size: 1. Harmarville,

2. do.

3. do.

4. do.

5. do.

6. do.

Cat. No. 61.4742 (cf) 149 mm. 90 mm. 51 mm. (largest)

“ “ 61.3667 ( 9 gravid) ... 148 “ 88 “ 56 “

“ “ 61.4742 (cf) 116 “ 68 “ 49 “

“ “ do. ( 9 gravid) . . . 109 “ 66 “ 40 “

“ “ do. (cf) 107 “ 64 “ 43 “

“ “ do. (cf) 93 “ 58 “ 31 “

Comparing these measurements with those of the creek-form, given above, it
is clearly seen that this form stands very close to the type, but grows larger, and
is relatively somewhat shorter and higher. It also is distinctly thinner in the shell.
It is in this form, that I have best recognized the sexual differences of the shell.

In addition there are other local forms in our region, and such are found
chiefly in the lakes of the glacial area in northwestern Pennsylvania. The fol-
lowing deserve special mention; but should not bear distinct names, since they are
only special reactions to special environment, and probably entirely inconstant.

Conneaut-lake-form A. Resembles in shape and the thin shell the form
gigantea, but is only half as large, and the epidermis has a peculiar light green to
pale brown color. I have only a few specimens, which I collected on gravelly
bottom on the north east shore of Conneaut Lake in from three to four feet ot
water.

Conneaut-lake-form B. Like form A in color (light green), but much more
elongated and subelliptical (not subovate). This form in shape looks almost like
A. marginata Say (= lacustris Lea), but traces of the beak-sculpture of grandis
are plainly seen. I have only a single specimen, which was collected by Mr. D.
Stewart, and I do not know particulars about the ecological conditions, except
that it comes from somewhat deeper water. L. 58 mm., H. 32 mm., D. 17 mm.

Conneauttee-lake-form. A much elongated form, like form B of Conneaut Lake,
with very thin shell, but epidermis darker, green to brownish black. As in form B
from Conneaut Lake, the beaks appear much anterior in consequence of the posterior
elongation of the shell. The typical (/randfs-sculpture of the beaks is well developed.

L. H. D.

Size: 1 115 mm. 55 mm- 39 mm.

2 76 “ 40 “ 22 “

3 50 “ 25 " 15 “

142

MEMOIES OF THE CAENEGIE MUSEUM.

I obtained five specimens of this form in the black muck of Conneauttee Lake,
Edinboro, Erie Co. In the outlet of the lake (Conneauttee Creek) the creek-
form of A. grandis was more or less typically developed.

Finally, it should be mentioned, that there are often specimens among typical
A. grandis, which are more swollen, thus approaching the var. footiana (to be dis-
cussed below) . Such specimens sometimes are rather frequent at certain localities,
chiefly so in the uppermost Shenango River, at Linesville, Crawford Co., and
these forms are truly transitional toward var. footiana. It also should be empha-
sized that the form in Conneaut Creek, at West Springfield, Erie Co. (tributary to
Lake Erie) is the typical creek-form of A. grandis.

Anodonta grandis behaves in Pennsylvania exactly as elsewhere. Over the
wide range of this species we have a great tendency to form local races and eco-
logical forms, which in many cases have received names of their own, without

Fig. 14.

• Anodonta grandis.

B Anodonta grandis footiana.

+ Anodonta ohiensis.

deserving them. I cannot go further into detail, since my material of the extra-
limital forms is not at all sufficient. But it is clearly seen that in Pennsylvania
as elsewhere this species is protean, accommodating itself to a variety of conditions,
and assuming different characters in shape and thicloiess of shell, in color, and in
size. It does not seem to reach in our state the extreme size recorded from other
parts (Baker, 1898a, p. 53, L. 171 mm.). However, one character generally

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143

remains constant, or varies only within narrow limits, that is the beak-sculpture,
which in well-preserved individuals is always double-looped and tubercular.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio-drainage:

Smiths Ferry, Beaver Co. (D. A. Atkinson).®*

Raccoon Creek, New Sheffield, Beaver Co.

Ice pond, Edgeworth, Allegheny Co. (G. H. Clapp) (forma gigantea).

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Brush Creek, Celia, Beaver Co.

Glade Run, Zeno, Butler Co.

Bonnie Brook, East Butler, Butler Co.

Wolf Creek, Grove City, Mercer Co.

Mahoning River, Hillsville, Lamence Co.

Neshannock Creek, Volant, Lawrence Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

Shenango River, Sharpsville, Shenango, and Jamestown, Mercer Co.; Linesville, Crawford Co.

A llegheny-drainage :

Pond, Harmarville, Allegheny Co. (forma gigantea).

Beaver Run, Delmont, Westmoreland Co.

Pond, Idlepark, Westmoreland Co. (D. A. Atkinson) (forma gigantea).

Crooked Creek, Rosston and Southbend, Armstrong Co.; Creekside, Indiana Co.

Cowanshannock Creek, Rural Valley, Armstrong Co. (N. M. Grier).

Sandy Creek, Sandy Lake, Mercer Co.

French Creek, Cochranton, Meadville, and Cambridge Springs, Crawford Co.

Sugar Creek, Cooperstown, Venango Co. (E. P. Spencer).

Conneaut Outlet, Conneautlake, Crawford Co.

Conneaut Lake, Crawford Co. (special forms, see above).

Cussewago Creek, Mosiertown, Crawford Co. (H. & L. Ellsworth).

Conneauttee Creek, Edinboro, Erie Co.

Conneauttee Lake, Edinboro, Erie Co. (special form, see above).

Leboeuf Creek, Waterford, Erie Co.

Brokenstraw Creek, Garland, Warren Co.

Connewango Creek, Russell, Warren Co.

Monongahela-drainage:

Monongahela River, Westmoreland Co. (G. A. Ehrmann).

Ten Mile Creek, Amity, Washington Co.

South Fork Ten Mile Creek, Waynesburg, Greene Co.

Dunkard Creek, Wiley and Mount Morris, Greene Co.

®* Not recorded upon the label whether from the Ohio or from Little Beaver Creek, probably from
the latter.

Rhoads (1899) does not give this species from Wampum, and H. H. Smith also apparently over-
looked it, since the specimen, upon which our record is founded, was discovered among duplicates of
Strophitus edentulus.

144

MEMOIRS OF THE CARNEGIE MUSEUM.

Lake Erie-drainage:

Conneaut Creek, West Springfield, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage :

Black Creek, Chili, Monroe Co., New York (R. H. Santens).

Creek at North Fairfield, Huron Co., Ohio (O. E. Jennings).®^

Ten Mile Creek, Lucas Co., Ohio (C. Goodrich).®'*

Miami and Erie Canal, Waterville, Lucas Co., Ohio (C. Goodrich).

Maumee River, Toledo, Lucas Co., Ohio (C. Goodrich).

St. Marys River, Rockford, Mercer Co., Ohio (C. Goodrich).

Beaver Creek, Williams Co., Ohio (C. Goodrich).

Raisin River, Adrian, Lenawee Co., Michigan (C. Goodrich).

Big and Little Whitefish Lake, Pierson, Montcalm Co., Michigan (Miss M. O’Malley). ®®

Okio-drainage :

Chautauqua Lake (R. Foerster): Bemus Point (D. R. Sumstine), Griffith Landing (Miss B. A. Ortmann),
Celeron (P. E. Nordgren), Chautauqua Co., New York.®®

Conotton Creek, New Hagerstown, Carroll Co., Ohio.

Tuscarawas River, Ohio (Holland collection).

Wolfe Creek, Washington Co., Ohio (W. F. Graham).

Redfield, Perry Co., Ohio (Juny collection).

Scioto River, Kenton, Hardin Co., Ohio (C. Goodrich).

Lewistown Reservoir, Logan Co., Ohio (C. Goodrich) (head of Big Miami). *

Wabash River, Fort Recovery, Mercer Co., Ohio (C. Goodrich).

Gravel pit, near Bluffton, Wells Co., Indiana (E. B. Williamson).®’

Winona Lake, Kosciusko Co., Indiana (E. B. Williamson) (Mrs. E. Courtney).®®

West Fork River, Lynch Mines, Harrison Co.; Lightburn and Weston, Lewis Co., West Virginia.

Little Kanawha River, Burnsville, Braxton Co., West Virginia.

North Fork Hughes River, Cornwallis, Ritchie Co., West Virginia.

Licldng River, Farmer, Rowan Co., Kentucky.

Fleming Creek, Pleasant Valley, Nicholas Co., Kentucky.

®® Intergrading toward var. footiana.

®* Intergrading toward var. footiana.

®® A form, compressed like typical grandis, but having a peculiar, light-colored epidermis.

®® I have many specimens, all representing a peculiar local race, greatly resembling A. benedictensis
Lea; small (maximum L. 90 mm.), and not quite as short as benedictensis. Most specimens much abraded
upon posterior slope. Color light green to light brown, but not blackish. Beak-sculpture of typical
grandis-charactcr. Call (1885) reports A. footiana from this lake, but my specimens are not this.

®’ Intergrading toward var. footiana.

®® A special form resembling gigantea in outline, but rather swollen, like footiana, and with the
beak-sculpture of typical grandis. Epidermis light greenish to brown. One specimen almost like speci-
mens from Tippecanoe Lake, recorded below under footiana. Call (1900) gives footiana from Eagle
(= Winona) Lake; Norris (1902) records grandis from Winona as well as Pike and Center Lakes, and
does not mention footiana.

ORTMANN: monograph of the naiades of PENNSYLVANIA,

145

Tennessee-drainage :

Mountain Fork Flint River, New Market, Madison Co., Alabama (H. E. Wheeler).®^

Western and southwestern localities:

Flat Creek, Sedalia, Pettis Co., Missouri (W. J. Utterback) (typical!).

Wakarusa River, Lawrence, Douglas Co., Kansas (R. L. Moodie) (typical!).

Sabine River, Logansport, De Soto Parish, Louisiana (L. S. Frierson).*''®

Lake Providence, East Carroll Parish, Louisiana (Juny collection).*®*

The Carnegie Museum possesses additional material, chiefly from southern and southwestern
localities, but these are local forms or races, with which I am not familiar, and so I do not discuss them
here. There are specimens at hand from the Alabama-drainage, in part labeled A. hallenbecki Lea by
Walker. They are very close to A. grandis, and if they are actually hallenbecki, this name should also
probably become a synonj^m of A. grandis.

Distribution and Ecology in Pennsylvania (See fig. 14) : As has been mentioned
above, the typical A. grandis is a form characteristic of smaller creeks. It decidedly
avoids the larger rivers, and is not found at all in the Ohio and Allegheny, and from
the Alonongahela only a single individual is on record. This holds good also
farther down the Ohio. In the tributaries, however, it is not rare, and becomes
most frequent in the smaller headwaters.

In these creeks it prefers quiet pools and eddies, often below riffles, where it
sticks in fine sand and mud. Occasionally it is found in riffles, but then probably
washed out of deeper and more quiet places. Yet it is to be seen that the typical
form by its somewhat stronger shell is adapted to a stronger flow of water, since
even in the quiet pools of small streams a stronger current may prevail during
periods of high water.

Under these conditions this species is found rather uniformly over western
Pennsylvania, atid goes eastwards to Warren, Indiana, and Westmoreland Cos.
But it is also capable of living in rather different surroundings, namely in ponds
and lakes. But we have seen that special forms develop under these conditions.
However, with the exception of a few glacial lakes, this environment is not often
found in western Pennsylvania, and there is a remarkable lack of ponds in the
southwestern part of the state. Thus the pond forms are rather scarce in our
region, as may be seen from the records given above, and we notice that they
often turn up in isolated ponds (Harmarville, Edgeworth, Idlepark), where there
is apparently no connection with streams inhabited by it. This has been observed

A large, thick-shelled form, corresponding wonderfully well with Lea’s original figure of gigantea
(Obs. II, 1838, PI. 1, fig. 1).

*®® A specimen absolutely indistinguishable from typical creek-forrns of Pennsylvania, possibly
not quite as thick-shelled as the average.

i®i Very much like the typical gigantea.

146

MEMOIRS OF THE CARNEGIE MUSEUM.

elsewhere, and its presence in such ponds can be explained only by the assumption,
that exceptional means of dispersal are available, presumably transport by aquatic
birds. This is also suggested by its distribution across the northwestern section
of our state, where it seems as if it had gone across country, not following the
streams. This, however, requires further investigation. It should also be pointed
out that it is very remarkable that these factors, if acting at all, did not act in an
eastward direction across the Alleghenian divide; and that this species is entirely
missing in the Atlantic-drainage, although it approaches the divide very closely
in Indiana and Westmoreland Cos. But this holds good only for Pennsylvania,
and is different in New York (see below).

General distribution: Type locality, Fox River of Wabash River, Indiana (Say).

According to Simpson (1900, p. 644),^°^ the range covers the “entire Missis-
sipjii system; the iqiper St. Lawrence-drainage ; Red River of the North; Lake
Winnipeg; Manitoba; southwest Texas” (he also with doubt mentions south-
eastern Pennsylvania, but this should be cancelled). This is certainly a correct
general statement. But the exact boundaries of this species are yet largely ill-
defined. In western Pennsylvania we apparently have part of the northeastern
boundary of the range, and it here coincides practically with the divide between the
interior basin and the Atlantic-drainage. However in New York state this is
different, for here this species passes over into the Atlantic-drainage of the Hudson
as well as the St. Lawrence River (see DeKay, 1843; Dewey,. 1856; Marshall,
1895; Baker, 18985). In part in its eastward advance, it seems to have used the
route of the Erie canal, reaching Albany County, and has here invaded the terri-
tory of the eastern A. cataracta. Its relation to this latter species in this region
(western and central New York) has never been closely studied, and on account
of the great resemblance of these two species it is very likely that they have often
been mistaken for each other. In this region A. grandis (generally known under
the name of A. lewisi) is found chiefly in streams and in some smaller lakes, and
it does not go into the large lakes. The same seems to be true all along the northern
boundary of its range, and, as soon as the great lakes are reached, it turns into the
var. footiana. In Pennsylvania, it goes up to the divide, and it is even in Conneaut
Creek, a tributary of Lake Erie; but it is not in the lake.

Generally in the region of the divide, intergrades are found between the two
varieties. Farther south grandis prevails, and finally is the only form in existence.

To Simpson belongs the credit of having reduced the number of species of Anodonta to reasonable
limits, and he has to a large extent worked out the synonymy of this species; but I hardly think that he
has gone far enough, and believe, that some “ species ” admitted by him, will finally fall as synonyms
under A. grandis.

ORTMANN: monograph of the naiades of PENNSYLVANIA,

147

In the Tennessee-drainage it is extremely rare, but certainly extends into northern
Alabama, It is also very rare in the Cumberland-drainage, but has been reported
by Wilson & Clark (1914) from ponds at Clarksville, Montgomery, Tennessee,
and from Stones River, in Rutherford Co,, Tennessee,

In the southwestern part of its range this species seems to become more
decidedly a pond- or lake-form. Call (1900) makes the statement that in Indiana
it is found in the ponds along the Ohio and Wabash, and that it occurs in the rivers
in the northwestern part of the state, that is to say, in smaller streams; this cor-
responds well with what we have observed in Pennsylvania, But west of the
Mississippi pond-forms prevail, and assume special shapes, which I cannot discuss
here, for they require much more detailed study (compare Wheeler, 1918, p, 121),

Anodonta grandis footiana (Lea) (1840),
Anodonta grandis footiana (Lea) Simpson, 1914, p, 422,

Plate X, figs, 3, 4,

Records from Pennsylvania:

Ortmann, 19096, p. 202,

Characters of variety: According to Baker (1898a, pp, 51 and 53), this variety
differs from Anodonta grandis by the greater inflation of the shell, chiefly in the
anterior and umbonal region, and by the finer beak-sculpture, which consists of
only four bars (while there are five in A. grandis).

This is the most concise statement of the differences of the two forms that
has ever been made. But with regard to the number of the bars in the beak-
sculpture it is to be remarked that in the light of my material, this is quite variable
(from four to six bars in footiana as well as grandis) , but with a stronger tendency
toward the smaller figure in footiana.

In other respects A. grandis footiana behaves much like A. grandis^ and varies
greatly in shape (more or less elongated), thickness, and color (from yellowish
and greenish to brownish and reddish) ; it lacks the dark green and blackish tints
found so often in grandis. Rays are generally entirely missing. The size is
inferior to that of grandis.

L. H. D.

Size: 1, Beach Pool G,, Cat. No. 61.4182 122 mm. 65 mm. 48

2. do. “ “ do 115 “ 57 “ 45

3. do. “ “ do 100 “ 52 “ 38

4. Beach Pool E. & F., Cat. No. 61.4181 ... 98 “ 53 “ 41

5. do. “ “ do. ... 95 “ 49 “ 37

6. “ Big Bend,” Cat. No. 61.4188 107 “ 67 “ 43

7. do. “ “ do , , , 93 “ 53 37

mm.

(largest at hand)

long form.

■short form.

4

148

MEMOIRS OF THE CARNEGIE MUSEUM.

Soft parts: As far as studied, identical with those of A. grandis, but gravid
females have never been found, and thus the glochidia and the breeding season
are unknown. However, the fact that the specimens, collected in May, June, and
July, were not gravid, indicates that the “interim” falls in these months, as is the
case in A. grandis, and thus probably the breeding season will be about the same.

Remarks: I regard this as a geographical race of A. grandis, which replaces
the latter in the St. Lawrence-drainage, and chiefly in the Great Lakes. It is,
however, not a “good species,” for south of the lakes, and passing somewhat beyond
the divide, into the Ohio-drainage, there is a region, where true intergrades are
found (in Pennsylvania, Ohio, Indiana, and Illinois). Some of these intergrades,
standing nearer to grandis, have been recorded under grandis (see above). And
further, even in Lake Erie, the form footiana does not always preserve its typical
characters, becoming sometimes more flattened (specimens from the lagoon at
Waldamer Park, and from ponds at mouth of Elk Creek, and occasionally else-
where), and having heavier beak-sculpture.

As regards the beak-sculpture, it is essentially of the grandis-type in so far
as it consists of four or five (sometimes six) bars, of which the third to the sixth
are distinctly double-looped, the loops separated by a sharp, re-entering sinus.
M ithin this sinus the bar is generally lower, sometimes almost effaced, so that it
represents a notch, on either side of which the anterior and posterior loop appear
elevated. However, the posterior loop of footiana, which has the same angular
and narrow shape as in grandis, is not so much thickened, and is not so distinctly
tuberculiform, and altogether the beak-sculpture is less heavy, finer, and sharper
than in grandis. The notch-like shape of the sinus, forming generally an inter-
ruption of the bar, shows that this form should be placed under A. grandis, although
by the delicacy of the sculpture it distinctly inclines toward A. cataracta.

In Lake Erie (in Pennsylvania), A. grandis footiana shows two extremes of
development of the shape of the shell, which are connected, however, by inter-
grades, and are, according to my observations, distinctly dependent on the eco-
logical conditions under which they live.

The most abundant form (among my material) is rather elongated, and closely
corresponds in the general outline to Lea’s maryattana (Obs. HI, 1842, PI. 20,
fig. 45), but it is not so excessively and abruptly swollen. In its typical develop-
ment this form is also characterized by a thin shell, and a pecuhar, rusty-brown
color, which is especially intense at and near the umbos, passing into brownish

103 In

some cases, the fifth, and chiefly the sixth bar (if this is present) have the sinus less sharp,
resembling only an undulation: this is a variation, which I have never distinctly observed in A. grandis.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

149

and yellowish brown toward the lower margin (alternating according to the growth-
periods). There is no, or very little, green present, and the rays upon the posterior
slope, if visible at all, are brownish black. According to my experience this form
is characteristic of the lagoons and beach-pools of Presque Isle, where it is quite
abundant, and very uniform in character.

The other form is less elongated, higher and shorter, sometimes squarish,
with a very straight hinge-line, having a sharp anterior and posterior angle. Its
shell is generally more solid, and its color is distinctly greenish, often rather light
green, but passing into light brown, rarely into rusty. Its growth-rests are distinct
and rather regular, and the rays upon the posterior slope are dark green. This is
the form prevailing in Presque Isle Bay, chiefly on its north-shore, so called “Big
Bend.” Some of these short, squarish forms resemble to a degree A. benedictensis
Lea (Obs. I, 1834, PI. 16, fig. 48; and DeKay, 1843, PL . 18, fig. 235), but I do not
think they should be called by this name,^'’^ for there are intergrades with the
elongated form of the beach-pools. These intergrades are frequently found
associated with the normal (quadrate) form, and do not have the quadrate shape,
but are more or less subovate, much resembling the original figure of Lea’s footiana
(Obs. Ill, 1842, PI. 20, fig. 44). The specimens collected in ponds at the mouth
of Elk Creek are all of this subovate form, with the typical footiana outline, and
at the extreme western end of Presque Isle Bay in quiet and rarely disturbed water
this ovate form is also present. In the lagoon at Waldamer Park I found speci-
mens, which are more elongated, and thus approach the beach-pool form, but they
have green (not rusty) epidermis.

It is evident that the squarish, thick-shelled form of the surf-beaten northern
shore of Presque Isle Bay passes gradually into the elongated, thin-shelled form of
the lagoons and beach-pools; and that it should be regarded as an ecological
variation of A. grandis footiana (with which it agrees in the inflated beaks), and
not as a form of benedictensis.

The various forms of footiana are distinguished from A. cataracta by the
swollen and inflated beaks. Although cataracta has sometimes a rather inflated
shell, the beaks are always low, and not so prominent as in footiana.

A. benedictensis comes originally from Lake Champlain, and has been reported from other lakes
in New York. The Lake Chautauqua-form of A. grandis, mentioned above, might be regarded as a
dwarf form of it, and also the form from Lake Erie (Sterki, 1907a, p. 394; Walker, 1913, p. 22). Mr.
Walker (according to a communication in a letter) considers the quadrate form and straight hinge-line
as the chief characteristics of his benedictensis, and admits that this differs from the true beriedictensis
in being more inflated. All this fits the present form.

150

MEMOIRS OF THE CARNEGIE MUSEUM.

Localities in Pennsylvania, represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, and beach-pools of Presque Isle, and lagoon at Waldamer Park, Erie,
Erie Co.

Pond at mouth of Elk Creek, Miles Grove (North Girard), Erie Co.

Other localities represented in the Carnegie Museum:

Lake Ontario, Braddock Bay, Manitou Beach, Monroe Co., New York (R. H. Santens).*®^

Lake Erie, Port Dover, Norfolk Co., Ontario, Canada (C. Goodrich).

Lake Erie, Sandusky Bay, Cedar Point, Erie Co., Ohio (0. E. Jennings) (normal).

Lake Erie, La Plaisance Bay, Monroe Co., Michigan (C. Goodrich) (young specimens).

Cedar Creek, Jerusalem Township, Lucas Co., Ohio (C. Goodrich).

Marsh, north of Toledo, Lucas Co., Ohio (C. Goodrich).

Miami and Erie Canal, Waterville, Lucas Co., Ohio (C. Goodrich).

Maumee River, Roche de Boeuf Rapids, Lucas Co., Ohio (C. Goodrich).^®®

Grand Reservoir, Mercer Co., Ohio (C. Goodrich).

Silver Lake, Clark Co., Oliio (Hartman collection).!®®

Tippecanoe Lake, Kosciusko Co., Indiana (E. B. Williamson).!®®

Lake Huron, Port Huron, St. Clair Co., Michigan (N. M. Grier).

Black Lake, Muskegon Co., Michigan (Exch. Alabama Museum).

Big Bass Lake and Au Sable Lakes, Lake Co., Michigan (C. Goodrich).

Douglas Lake, Cheboygan Co., Michigan (H. B. Baker).

Muskoka Lake, Bala, Muskoka Co., Ontario, Canada (A. S. Daggette).

Lake Helen, Nipigon, Ontario, Canada (0. E. Jennings).

Lake Superior, Nipigon Straits, St. Ignace Island, Ontario, Canada (0. E. Jennings).

Blackwater River, Jellicoe, Ontario, Canada (0. E. Jennings) (Tributary of Lake Nipigon).

Long Lake, Longuelac, Ontario, Canada (0. E. Jennings).

In 1914, Dr. Jennings collected a form in Lake Nipigon (Orient Bay), which, in shape and beak-
sculpture, is a true A. grandis, but has a peculiar rough epidermis. I have not yet been able to identify
this.

Distribution and Ecology (See fig. 14) : Type locality, Winnebago, Winnebago
Co., Wisconsin (Lea).

In Pennsylvania this form is restricted to Lake Erie, and although individuals
resembling it have been found in the uppermost Shenango, they were associated

!®® An interesting set of sixteen specimens, containing individuals inflated like typical footiana,
some much more flattened, almost like typical grandis, with intermediate stages between these. Outline
of the shell elongated-ovate, shorter or longer, like the creek-form of grandis, beak-sculpture more like
grandis. Color light green to gray or brownish green. These are truly intergrades.

!®® A special (river-) form, heavy shelled, greenish, with rather heavy beak-sculpture. Considerably
inflated at the beaks. A link in the chain of intergrades.

!®! Inflated like footiana, but in outline resembling the gigantea type.

!®® A typical elongate-subovate, greenish footiana. The specimen was labeled as given, but I can-
not find a Silver Lake in Clark Co., but there are places of this name in Logan and Summit Cos.

!®® Wabash-drainage. A peculiar large form, resembling the form of grandis gigantea from Winona
Lake, but more swollen, and with the beak-sculpture of footiana.

ORTMANN: monograph op the naiades op PENNSYLVANIA.

151

with typical grandis, and should be classed with them. In Lake' Erie, this form
is found, as has been stated, under various ecological conditions, but not in the
open lake. It is present on the north shore of Presque Isle Bay, where there is,
at times, a considerable surf, and lives here in from one to five feet of water. It
has also been brought up by the “sand-sucker” from a depth of ten to fifteen feet.
It prefers a bottom of pure sand or fine gravel, either bare, or covered with a growth
of rushes {Juncus americanus). Many specimens on the north shore of the bay
are much abraded, and apparently worn by the surf. Toward the western end
of the bay, it is found upon the sandy “flats,” among rushes, and at the extreme
western end on a bottom, which consists of a mixture of sand and mud; in this
section, the water is rather quiet, there is surf only during easterly winds, and
this is broken and checked by the Juncus americanus formation, which runs far
out into the bay. Further, this species in a special form inhabits the numerous
lagoons and beach-pools of Presque Isle, which are rather shallow, and have a
bottom of fine sand, often covered with more or less fine vegetable mud. It occurs
also in the large and deep lagoon at Waldamer Park (base of Presque Isle), and I
also found it in a pond cut off from the mouth of Elk Creek, on sandy-gravelly
bottom. In Pennsylvania, this is distinctly a lake- and pond-form.

Lake Erie and the smaller lakes of the state of Michigan seem to be the metrop-
olis of this form. The largest number of locality records are at hand from Michigan,
due to the fine work of Walker (1898, see map of distribution, PI. 1). Northwards
it goes here as far as Chippewa Co. (Winslow, 1917) and Marquette Co. (South
of Lake Superior), and Lake Gogebic, Ontonagon Co. (Ruthven). It is also in
the lakes and bays of Isle Royale in Lake Superior (Walker, 1909, p. 294), and the
Carnegie Museum possesses it from the region of Nipigon, on the North shore of
Lake Superior. The most extreme locality in a northwestern direction is Souris
River, Manitoba (Hudson Bay-drainage) (Dawson, 1875).

From Wisconsin it is known only from the type locality (Winnebago). South-
ward, it crosses over into the Mississippi-drainage in northern Ohio.(Sterki, 1907a),
northern Indiana (Call, 1896a and 1900),^^° and northern Illinois (Cook, Lake,
Kane, McHenry Cos.) (See Baker, 1906, p. 73).

From Lake Erie eastwards and northeastwards, we have a number of records
in western New York (Marshall, 1895), as far as Ottawa River, Canada (Call,
1885). However, some of these are doubtful, as for instance. Call’s record from
Chautauqua Lake, according to him the only locality in the Mississippi basin.

As to grandis and footiana in Winona Lake (See footnote 98 on p. 144). Our specimens from this
lake are nearer grandis. The form from Tippecanoe Lake in our collection is more like footiana; Wilson
& Clark (1912a, p. 47) call this form: grandis, “ of a more inflated type.”

152

MEMOIRS OF THE CARNEGIE MUSEUM.

The form found in this lake, represented in the Carnegie Museum by numerous
specimens, is a small race of A. grandis, possibly to be placed with henedictensis,
but not with footiana.

A. grandis footiana belongs preeminently to the St. Lawrence system, and
has a wide range in it. It crosses over into the Mississippi as well as the Hudson
Bay drainages, but only very slightly, and in the former it intergrades with A.
grandis.

Anodonta cataracta Say (1817).^^^

Anodonta cataracta Say. Simpson, 1914, p. 386.

Plate X, fig. 5; Plate XI, fig. 1.

Records from Pennsylvania:

Haldeman, 1844 (Lancaster Co.).

Gabb, 1861 (Mill Creek, Milltown; Wissahickon Creek and Schuylkill River, Philadelphia, Germantown
and League Island, Philadelphia Co.).

Lea, 1867 (Schuylkill, Delaware River, and League Island, Philadelphia) (tryoni).

Bruckhart, 1869 (Lancaster Co.).

Hartman & Michener, 1874 (Schuylkill River, Chester Co.).

Pilsbry, 1894 (York Furnace, York Co.).“^

Schick, 1895 (Delaware and Schuylkill Rivers, Fairmont Park, League Island, Philadelphia; German-
town, and Canal at Manayunk, Philadelphia Co.; Munckinipattus Creek, Glenolden, Delaware
Co.).

Marshall, 1895 (Tributaries of Genesee River, Potter Co.;“® Juniata River).

Ortmann, 19096, p. 205.

Caffrey, 1911 (Lehigh Canal and ponds, Bethlehem, Northampton Co.).

Characters of the shell: This species is very close to A. grandis, and the only
reliable difference is in the beak-sculpture, and even this is a difference rather of
degree of development than of type. The general character of the beak-sculpture
is the same in both species (double-looped), but while in A. grandis the two loops
are sharply separated by a sinus, which is generally depressed (notch-like), and
while the posterior loop is tuberculiform, in A. cataracta the bars of the beak-
sculpture,'^ which are also slightly more numerous (five to seven) are uniform in
elevation, and in the sinus they are not appreciably lower than in front and behind
of it, so that no tubercles are formed. These differences have properly been pointed
out by Marshall (1890, p. 188, 189). In the light of my material I have only to
add, that the first two bars are simplj^ concentric, and that those following are

Not 1816, see p. 9.

Pilsbry’s (p. 30) “ A. subcylindrica Lea ” is this species. I have seen the specimens so labeled
in the Philadelphia Academy of Natural Sciences.

This locality should be confirmed.

ORTMAN’N: monograph of the naiades of PENNSYLVANIA.

153

double-looped, but in the last (sixth and seventh) the sinus is often not sharp, but
represented merely by a gentle undulation. There is a good deal of variation,
chiefly in the number of bars, but also in the sharpness of the sinus, which may be
acute or more or less blunt.

In other characters the shells of the two species are much alike, but as a rule
A. cataracta inclines more toward the elongated shape, although it also possesses
comparatively short and high forms. In the elongated pond-forms, another
difference is noted in the position of the posterior end, which is more elevated
above the base-line, so that the lower margin of the shell is more strongly convex,
its posterior part running more decidedly upwards. But this character is not
observed in the creek-forms. Also the posterior “wing” is generally more distinct
in A. cataracta, although often very low.

On the average the color of A, cataracta is brighter green than in A. grandis.
It is light to dark green, rarely shading into brownish and blackish, often very
brilliant, with concentric lighter and darker bands, and the dark green rays are
also more distinct upon the disk; though there are specimens which do not show
any rays. The broad rays of the posterior slope are often strongly developed
and broadened, so as to render the whole posterior slope blackish green.

In this species as in its allies, the female sex is sometimes indicated by a swelling
of the valves posteriorly to the middle, but not all females exhibit this character.

L.

H.

D.

1. Flinton, Cat. No. 61.3637 (largest on hand) . .

. . 128 min. 72 mm.

45 mm.

2. do. “ “ 61.3636

. .125

“ 65 “

44 “

- pond-form

3. Lockhaven, Cat. No. 61.4173 ( 9 gravid) . . . .

. .112

“ 56 “

42 “

4. Tioga, Cat. No. 61.4171 (9 gravid)

.. 85

“ 46 “

27 “ ■

5. Avondale, Cat. No. 61.4167

.. 86

u ^2 “

31 “

- creek-form

6. York Haven, Cat. No. 61.4727 (c^)

.. 84

« 47 »

28 “

7. Schuylkill, Philadelphia, Cat. No. 61.816. . . .

.. 87

“ 50 “

33 “

iryom'-type

These measurements do not express the maximum size, and considerably
larger specimens are on record.

Soft parts practically identical with A. grandis (See Ortmann, 1912, p. 293).
Glochidia figured by Lefevre & Curtis, 1910, p. 97, fig. C, and 1912, p. 146, fig. C.

Breeding season: Conner says (1907 and 1909) that this species is gravid from
September to May, and that the interim falls in June, July, and August. I have
collected gravid females on July 23, 1908; August 19, 1909; Aug. 21, 1908; Aug.
24, 1909; Sept. 5, 1909; March 19, 1911; April 24, 1909.

Of these dates, July 23 is surely an exceptional case (only one specimen was
gravid among a large number). The dates in August mark the beginning of the

154

MEMOIRS OF THE CARNEGIE MUSEUM.

breeding season, and are earlier than Conner’s records. Some of the specimens
collected on April 24 were found discharging. This would establish the breeding
season as lasting from August to May, a condition, which is much like that seen in
A. grandis. It seems, that the glochidia are discharged rather early, for I never
found any charged females in Alay.

Remarks: There still might be some lingering doubt, as to whether A. cataracta
is specifically distinct from A. grandis. However, I have never seen any specimens,
which might be called intergrades, and wherever I have examined the beak-sculp-
ture, it has always been possible to distinguish these two species by this character.
Nevertheless they are closely allied, and there is no question that A. cataracta is
the eastern representative of the western A. grandis. The two forms are geo-
graphically well separated. It is true that in New York the ranges do overlap,
but I think this is due to a secondary, postglacial expansion of the two species
into the glacial area. During glacial times, they undoubtedly were completely
separated (Ortmann, 1913, pp. 325, 363-365). Walker’s opinion, expressed
incidentally, that cataracta is closely allied to the European A. cygnea, and might
be a co-immigrant with Margaritana margaritifera from Europe, cannot be main-
tained. Compare also Latchford (1914, p. 10).

A rather strong argument for the specific distinctness of A. grandis and cataracta
is, as I believe, their different behavior or reaction to environment. Both possess a
creek- and a pond-forjn, but while the reaction to the pond-environment in A.
grandis is to develop a large, high and comparatively short shell {A. grandis gi-
gantea), in A. cataracta a more elongated shell is the result, with a “rostrate”
posterior end. This latter characteristic shape of the shell is never found distinctly
in A. grandis. On the other hand, the creek-form of A. grandis is subovate, with
the anterior end broad, and the posterior narrowed, while the creek-form of A.
cataracta is rather subelliptical, with the anterior and posterior ends generally
rather equally narrowed.

The short and high type of shell, so common in A. grandis, is much rarer in
A. cataracta, but it is present. Lea’s “species” williamsi, virgulata, and tryoni
represent this. They may be described as follows.

A. williamsi. {Type locality Lower Potomac.) High and short, with better de-
veloped posterior wing, so that the upper and lower margins of the shell become
subparallel. Sometimes the posterior wing may be so high that the posterior
part of the shell appears higher than the anterior (ovate, with anterior end nar-
rower) .

A. tryoni. Closely resembling A. williamsi, but with the posterior wing not

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155

so high, so that the shape becomes short-ovate, with the posterior end nar-
rower.

A. virgulata. {Type locality North Carolina.) In shape intermediate between
A. williamsi and A. tryoni, but with brighter coloration and more distinct ra3^s.
This is, according to Simpson (1914, p. 388) a southern form of A. cataracta.

Such short and high shells seem to be rather scarce in Pennsjdvania, but the
tryoni-type occurs in the larger rivers (Delaware and Schu^dkill), and might be
called characteristic of this environment.

The thickness of the shell is also variable in A. cataracta; the pond-forms
being generally thinner, while the creek-forms are thicker. A. cataracta is also
quite variable in the convexity of the valves; and there are more compressed and
more inflated shells, both in ponds and creeks. The inflation, if present, is re-
stricted to the disk, and does not extend to the umbos, and forms having inflated
beaks, like A. grandis footiana, are not known among the variations of A. cataracta,
except possibly in the South, among the virgulata-type, in which there seems to
be a tendency in this direction.

The various forms of A. cataracta, described above, are all connected by inter-
grades, and the}^ seem to be special reactions to special environmental conditions,
although we are not in all cases sure what are the essential features of the environ-
ment, which are active. In the Schu^dkill Canal I found, for instance, shells
resembling the creek-form (but thin-shelled), where we should rather expect the
pond-form. But here the bottom of the canal was somewhat stony, with very
little mud, and it is quite possible, that in this case the character of the bottom was
all important. Specimens from Echo Lake, Monroe Co., more closely resemble
the creek-form. Since these various forms turn up all over the range of the species,
they are to be regarded simply as variations, not as varieties. The idea, that they
may be species is entirely excluded.

I should add here a few words about a peculiar, distorted form, received from
Mr. C. H. Conner from the Delaware River, Newbold, Gloucester Co., New
Jersey. There are five specimens, resembling in general characters the tryoni-
type, but all possessing a peculiar radial rib, running from the beaks toward the
lower margin of the shell, and in front of the rib the shell is somewhat compressed.
It seems as if the growth of the shell in its anterior part had been checked or re-
tarded. The specimens look decidedly like cripples, and I take them for such,
yet it is remarkable that a number of specimens showing the same deformation
should be found at the same locality.

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MEMOIRS OF THE CARNEGIE MUSEUM.

Localities in Pennsylvania represented in the Carnegie Museum:

Delaware-drainage :

Delaware River, Penn’s Manor, Bucks Co. (young, approaching tryoni).

White Clay Creek, Avondale, Chester Co. (typical creek-form).

Crum Creek, Delaware Co. (Hartman collection) (creek-form, but thin).

Schuylkill Canal, Manayunk, Philadelphia Co. (thin-shelled creek-form).

Schuylkill River, Philadelphia (Hartman collection) (labeled tryoni, and representing this type).
Wissahickon Creek, Roxboro, Philadelphia Co. (creek-form).

Little Neshaminy Creek, Grenoble, Bucks Co. (creek-form).

In a dry pond of Mahoning Creek, Lehighton, Carbon Co. (pond-form).

Echo Lake, Monroe Co. (G. H. Clapp, donor).

Susquehanna-drainage :

Conewago Creek, York Haven, York Co. (creek-form).

Penn Canal, Selinsgrove, Snyder Co. (lake-form).

Chemung River, South Waverly, Bradford Co. (creek-form, remarkably light green, found in riffles).
Mill-race of Crooked Creek, Tioga, Tioga Co. (typical creek-form).

Ponds near paper-mill, Lockhaven, Clinton Co. (all typical pond-form).

Beaver Dam Creek, Elinton, Cambria Co. (D. A. Atkinson) (typical pond-form; from an old dam in
the creek; some of the younger ones incline toward the creek-form).

Potomac-drainage :

Conococheague Creek, Greencastle, Franklin Co. (creek-form, unusually thick-shelled; from riffles in
water heavily charged with lime); Scotland, Franklin Co. (creek-form, dwarfed).

Great Tonoloway Creek, Thompson Township, Fulton Co. (creek-form).

Localities in Pennsylvania represented in the Philadelphia Academy of Natural
Sciences:

Eastwicks Park, Grays Ferry, Philadelphia (S. R. Roberts).

Wister Dam, Germantown, Philadelphia Co. (W. Stone).

Oxford Dam, Guinea Creek,*i^ Woodbourne, Bucks Co. (H. W. Fowler).

Porters Lake, Pike Co. (S. N. Rhoads) (headwaters of Bushkill Creek, to Delaware).

Harvey Lake, Luzerne Co. (C. H. Conner) (W. Stone) (Harvey Creek, to North Branch Susquehanna).
On New Jersey side of Delaware:

Pond near Delaware River, Delanco, Burlington Co., New Jersey (H. A. Pilsbry).

Other localities in the Carnegie Museum:

Monmouth, Kennebec Co., Maine (Woods collection) (typical creek-form).

Charles River, Cambridge, Middlesex Co., Massachusetts (Mrs. L. D. Thompson) (creek-form).

Lake Ontario, Braddock Bay, Manitou Beach, Monroe Co., New York (R. H. Santens).“®

A dead and broken shell was seen, but not taken, in the Juniata River (Raystown Branch) at
Ardenheim, Huntingdon Co.

No such name is on the topographical map; the creek at Woodbourne is a branch of Mill Creek.
One specimen, collected together with specimens of A. grandis footiana, mentioned above. It
has the typical shape of the creek-form of A. cataracta, and differs from the others in the beaks, which
are not inflated. Beak-sculpture poorly preserved, but apparently of the cataracta-tjpe.

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157

Backwater of Hudson River, Milton, Ulster Co., New York (H. H. Smith) (j^oung, intergrading from
pond-form to tryoni-iy^^e) .

Binghamton, Broome Co., New York (H. H. Smith).“^

Culvert Pond, Sussex Co., New Jersey (J. F. L. Raeschen) (creek-form, but thin-shelled).
Delaware-Raritan Canal, Aquaeduct, near Princeton, Mercer Co., New Jersey (young, h'i/o?n‘-type) .
Delaware River, Fish House and North Cramer Hill, Camden Co., New Jersey (from both places the
creek-form); Newbold, Gloucester Co., New Jersey (C. H. Conner) (cripples).

Potomac River, Washington, D. C. (Hartman collection) (iryonf-type).

South Branch Potomac River, Romney, Hampshire Co., West Virginia (chiefly creek-form, but some,
from quiet pools, inclining to pond-form).

South River (upper Shenandoah), Waynesboro, Augusta Co., Virginia (thin-shelled creek-form from a
quiet pool).

Paper-mill, Salem, Forsyth Co., North Carolina (Holland collection) {virgulata-iyY)Q) .

Ashburn Creek, Mecklenburg Co., North Carolina (Hartman collection) (virgulata-type) .

Distribution and Ecology in Pennsylvania (See fig. 15) ; In Pennsylvania this
species belongs to the Atlantic-drainage. It has indeed been reported by Marshall

• Anodonta cataracta. -f Anodonta miplicata.

(1895) from the headwaters of the Genesee River in Potter Co., but this requires
confirmation. When I collected in the Genesee River and Cryder Creek, at Gen-
esee, Pennsylvania, I did not find any Anodontas.

As the records show, A. cataracta is found in the drainages of all three Atlantic
rivers, the Delaware, Susquehanna, and Potomac. It apparently is most abundant
in the lowlands and on the Piedmont Plateau, but goes up considerably towards

The label says: “ Chemung River,” but Binghamton is not on this River; the specimen has
all the earmarks of a pond-shell.

158

MEMOIRS OF THE CARNEGIE MUSEUM.

and into the mountains, farthest in the Susquehanna, where it ascends to Cambria
Co., west of the Allegheny Front. Here it is closest to the divide, and it is inter-
esting to note, that on the other side in the Ohio-drainage of Indiana and West-
moreland Counties, the creek- and pond-forms of A. grandis turn up.

I collected the pond-form in rather small (artificial) ponds, with very muddy
bottoms, and also, but more rarely, in quiet pools in creeks. I obtained the
creek-form most commonly, but it is rather erratic in distribution. It seems to
avoid the larger rivers (although found under special conditions in the estuary of
the Delaware). It favors smaller rivers (Chemung) and creeks. Here it lives on
gravelly bottoms, in more or less strongly flowing water (even in riffles), or in more
quiet pools in gravel, sand, or mud. The short and high form (^ryonf-type) seems
to prefer large rivers with muddy bottoms, but I have not observed it often enough
to express a final opinion.

Probably this species has a more general distribution in eastern Pennsylvania,
and the pond-form might especially be found in many other ponds (and lakes) :
these localities, however, are often not easily accessible, and it is hard to locate
this shell in them.

General distribution: Type locality, not given by Say, but probably is eastern
Pennsylvania.

Simpson (1900) reports this species from “Lower St. Lawrence-drainage;
streams draining into the Atlantic south to North Carolina.”

In the St. Lawrence-drainage, it has been reported from Lake Ontario (Hamil-
ton Bay, Marshall, 1895, and Toronto, Latchford, 1914), and this is confirmed by a
specimen in the Carnegie Museum. Farther down it is known from various points
along the St. Lawrence, and also from the Lake Ontario-drainage in western New
York. But there is some doubt whether all the records given by Marshall (1895)
from this section actually refer to this species. At any rate two of these records:
Buffalo Creek in Erie Co. (to Lake Erie), and Ischua Creek, Cattaraugus Co. (to
Allegheny River) appear to me veiy doubtful, and should be confirmed. In this
region, indeed, it seems, that the ranges of A. grandis and A. cataracta overlap,
and on account of the great resemblance of these two species, it is quite possible,
that some of the identifications are not correct. This question deserves special
study.

On the Atlantic side this species goes northward to Maine, as is shown by
one of our localities, and it has been reported by Jackson (1908) North Haven
Island, Knox Co. and Lermond (1909) Cumberland Co. Walker & Coolidge
(1908) give it from Keene, Cheshire Co., New Hampshire, Adams (1842) from

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159

Middlebuiy, Addison Co., Vermont, and Alarshall (1895) from Benson, Rutland
Co., Vermont. It is also present in Rhode Island (Carpenter, 1890), and is frequent
in the western and central parts of Massachusetts (Gould-Binney, 1870). Thence
southward it becomes rather common, but south of Pennsylvania records become
scarce. No positive localities are at hand from Maryland. From Delaware it
has been mentioned by Rhoads (1904), and the localities from Virginia and North
Carolina are few. I have found it in the Potomac-drainage in West Virginia and
Virginia. According to Simpson it goes only to North Carolina, but there are
old records given by Lea for his virgulata and williamsi from Georgia, so that the
southern boundary of this species becomes obscure; in addition a number of
closely allied forms or species have been described from the southern states, the
standing of which remains to be investigated.

We must regard A. cataracta as a form characteristic of the northern section
of the Atlantic Coastal Plain and the Piedmont Plateau, from which center it has
spread more or less toward the Allegheny Mountains, but without crossing the
divide; and from which it has widely spread, probably in Postglacial times, to the
northeast and north, and also slightly northwestwards, reaching New England
and the lower St. Lawrence basin, where it has met the eastward expansion of the
western A. grandis (See Ortmann, 1913a).

Anodonta implicata Say (1829).

Anodonta implicata Say, Simpson, 1914, p. 391.

Plate XI, figs. 2, 3.

Records Jroin Pennsylvania :

Lea, 1838 (Schuylkill River, Philadelphia) (as A. newtonensis).

Gabb, 1861 (League Island, Philadelphia).

Hartman & Michener, 1874 (Schuylkill River, Chester Co.).

Marshall, 1895 (Pliiladelphia).

Ortmann, 19096, p. 206.

Characters of the shell: Large, elongated, similar in shape to the pond-form
of A. cataracta, but with the posterior end not so elevated; rather thick-shelled,
and noticeably thickened along the lower anterior margin (from the middle for-
wards). Nacre more or less reddish, or salmon-color.

. It is also said that the shell is more inflated (subcylindrical) than in A. catar-
acta, and that the color of the epidermis is lighter (yellowish) and has no rays.
This, however, probably is not always the case. Marshall (1890) says, that the
beak-sculpture is also different, with the sinus less developed, being gentle in the

160

MEMOIRS OF THE CARNEGIE MUSEUM.

earlier, and missing in the later bars. This holds good in at least one of the speci-
mens before me. The rough epidermis is also given as a character, as distinguished
from the shining cataracta, but this is not confirmed by my material.

L. H. D.

Size: 1. lardley, Cat. No. 61.36,39 75 mm. 41 mm. 26 mm.

2- do. “ “ do 74 “ 40.5 “ 25 “

Ihese are the only specimens of considerable size collected by myself, and
they are hardly more than half the maximum size attained by this form.

Sojt parts and glochidia unknown.

Breeding-season: According to Conner (1909) from September to May (like
cataracta). Lefevre & Curtis (1912, p. 141) enumerate this species under the forms
with long breeding-season.

Remarks: This is a form the taxonomic standing of which is as yet obscure
and I myself have not been able to form a clear conception of it. According to
previous authors, the characters given above are essential, but they apply only
to the adult shell, and I am unable to say much about the young shell. The
smallest shells I have seen are those I collected in the Delaware at Shawnee (Length :
58, 46, 41, and 39 mm.). Of these the largest shows traces of the thickening at
the lower anterior margin, while the smaller ones do not. These shells have the
epidermis rather yellowish, and the nacre has a good deal of salmon-color, and
therefore I think they actually represent this species. All other shells I have
seen are larger, and show the character of the thickening of the margins well-
developed. The best account of this shell has been given by Gould (Binney,
1870, p. 180), but even here it is admitted that the young shells are difficult to
distinguish from A. fluviatilis (= cataracta).

The Carnegie Museum possesses only eight specimens, to which I should apply
the name implicata. One (without locality) is from the Hartman collection, and
is certainly this form. It is large (L. 122 mm.), elongate (H. 67), and swollen
(D. 56). Its epidermis is shining (not rough) and of a yellowish-olive to light
brown color, without any rays. The beak-sculpture is as described by Marshall.
The valves also show the characteristic thickening along the anterior lower margin,
and the nacre is pale pinkish. Another specimen was sent to me by C. H. Conner
(Timber Creek, Gloucester, New Jersey, just south of Newton Creek, Camden
Co., the type locality of Lea’s newtonensis) . It is 103 mm. long, 49 high, and 41
in diameter. Thus it is rather elongated, swollen, and subcylindrical. The
epidermis is blackish brown (not pure black as in the figure of newtonensis), the
beaks are eroded, and the beak-sculpture gone. The thickening of the margin
is present, and the nacre is pinkish or salmon.

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161

In addition, I have collected two specimens in the Delaware River in Bucks
Co., which are the specimens of which the measurements are given above, and
to them must be added the four young ones, mentioned above, taken at Shawnee,
Monroe Co. All these are much smaller, and are not at all swollen nor subcyl-
indrical. The color of the epidermis is greenish with light brown or yellowish
concentric bands, without distinct rays, or else nearly uniformly light brown,
darker behind. The larger ones show the thickening of the shell, and all have a
more or less distinct tint of salmon in the nacre.

In the collections of the Philadelphia Academy of Natural Sciences I have
seen comparatively few specimens, hardly more than a dozen. The thickening of
the margin of the shell is always present, often also the pinkish color of the nacre
and the inflation of the valves. The color of the epidermis ranges from greenish
yellow to blackish.

Localities represented in the Carnegie Museum:

Delaware River, Yardley, Bucks Co.; Shawnee, Monroe Co., Pennsylvania.

Timber Creek, Newbold, Gloucester Co., New Jersey (C. H. Conner).

Localities represented in the Philadelphia Academy:

Schuylkill River (I. Lea).

Thorps Mill-Pond, Branchton, Philadelphia (S. R. Roberts).

Delaware River, Torresdale, Philadelphia Co., Pennsylvania (S. R. Jacob).

Channel of Delaware River, about 600 feet off New Jersey shore, in 35-45 feet depth, muddy bottom,
“ East of North end of Little Tinicum Island ” (L. Woolman).ii®

Distribution and Ecology (See fig. 15) : Type locality, Pond, Danvers, Essex
Co., Massachusetts (Say).

Simpson (1900) gives the range of this species as: “St. Lawrence-drainage ;
north to Lake Winnipeg; south in streams flowing into the Atlantic to Virginia.”
I am unable to substantiate this, and the localities on record only in part confirm
Simpson’s statement. The question also arises, whether all the citations recorded
actually refer to this species. At all events, I seriously question the record “Lake
Erie, Port Dover, Canada” (Marshall, 1895). South of New Jersey and Penn-
sylvania I know of no reliable records whatever.

In Pennsylvania this species has only been found in the neighborhood of
Philadelphia, in the Schuylkill and Delaware Rivers, and in ponds. It occurs also
on the opposite side of the Delaware in New Jersey. My own localities are farther
up the Delaware. Not a single specimen has ever turned up in the Susquehanna

Little Tinicum Island has no north end; probably what is meant is “ south of east end.” The
locality is off Billingsport, Gloucester Co., New Jersey.

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MEMOIRS OF THE CARNEGIE MUSEUM.

and Potomac drainages. Altogether it seems to me, that this form is more northern
in its distribution, and the majority of localities are from New York, Connecticut,
and Massachusetts, in which direction it goes as far as Maine (Lermond, 1909).
Generally it is reported as living in ponds, but sometimes also in rivers. The
locality in the deep channel of the Delaware (35 to 45 feet depth), given above, is
quite interesting and possibly suggestive. Further studies on this species are verj^
desirable. (See Ortmann, 1913a, p. 363.)

Anodonta ohiensis Rafinesque (1820).

Anodonta imbecillis Say. Simpson, 1914, p. 395; Lastena ohiensis (Rafinesque)

Utterback, 1916, p. 109.^^®

Plate XI, fig. 4.

Records from Pennsylvania :

Ortmann, 19095, p. 195 and 202.

Characters of the shell: Shell small, at the utmost of medium size, very thin.
Outline subelliptical, rather elongate, rounded anteriorly, narrowed and somewhat
pointed behind. I^Pwer margin gently convex, less so in the posterior part, some-
times almost straight in the middle. Upper margin straight, forming a more or
less distinct angle with the obliquely descending posterior margin, and in general
this angle is elevated to form a wing, which is rarely entirely obliterated. Beaks
depressed, and not at all elevated above the hinge-line. In young shells the beaks
slope down obliquely (roof-like) from the hinge-line; in older, more swollen shells,
they are nearly or entirely horizontal. Beaks placed anterior to the middle of
the shell. Beak-sculpture fine, weak, and irregular, consisting of four to six
subconcentric ridges, of which the later ones have a faint sinuation, which is only
wavy, and does not form an angle. Often this sinus is represented only by an
interruption of the bar.

Shell of young specimens very slightly swollen, or even compressed; in older
ones it becomes more inflated, and assumes in extreme cases an almost subcylindrical
shape. Posterior ridge very indistinct, rounded; greatest diameter of the shell
about in the middle. Posterior slope more or less compressed and elevated into
the wing of the upper posterior angle.

Epidermis lighter or darker green, often beautifully grass-green, grayish,
brownish, or yellowish near the beaks, with concentric, darker growth-rests, and

I object to the use of the generic name Lastena for this species. It is not the type of Rafinesque’s
subgenus Lastena, and a type for this genus was not designated until Simpson (1900, p. 654), selected
lata Rafinesque for it. It might possibly be desirable to generically separate this species from the other
species treated in this paper under Anodonta, but in this case, a new name should be found.

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163

with dark green, more or less distinct, rays on the whole disk. Two or three dark
green to blackish rays generally well developed upon the posterior slope.

Hinge edentulous, forming a straight line, or with a faint indication of a wave
under the beaks. Beak-cavity shallow. Dorsal muscle-scars in the beak-cavity.
Adductor muscle-scars indistinct. Nacre blueish white, silvery, iridescent, some-
times slightly cream-color toward the beak-cavities.

This species being a hermaphrodite does not show any sexual differences in

the shell. ^

L. H. D.

Size: 1. Erie, Cat. No. 61.3291 83 mm. 41 mm. 28 mm.

2. Waterford, Cat. No. 61.4178 67 “ 32 “ 24 “

3. do. “ “ do 56 “ 27 “ 16 “

4. Edinboro, Cat. No. 61.3290 48 “ 25 “ 12 “

The largest specimen represents about the maximum size attained by this
species. Generally it remains considerably smaller (Length between 50 and
60 mm.).

Soft parts (See Ortmann, 1912, p. 291). Glochidia figured by Lea (Obs. VI,
1858, PI. 5, fig. 36); Ortmann (1911, PL 89, fig. 13); Surber (1912, PI. 1, fig. 2),
Surber’s measurements are: 0.31 X 0.29 (longer than high), while I gave: 0.30 X
0.31 mm. (higher than long).

Breeding-season: My dates for gravid females are: June 2, 1908 (eggs);
July 12, 1910 (discharging); Sept. 14, 1909 (eggs and glochidia); May 21, 1908
(glochidia and discharging); May 22, 1909 (glochidia and discharging); May
24, 1909. Surber (1912, p. 7) gives March, June, July, August, and September as
months when gravid individuals are present. From Arkansas I have gravid
specimens collected by Wheeler on February 20, 1913; June 14, 1911; July 17,
1911; and in November, 1911.

This is undoubtedly a hradytictic form, and according to the above dates, it
apparently breeds “all the year round,” i.e., the succeeding breeding seasons
overlap in June and July, but probably not in the same individual.

Remarks: A well-marked species, which is easily recognized by its shape and
color. As far as my material permits a conclusion, it varies very little, and the
variations are restricted to the proportional length of the shell and its degree of
obesity; the latter of which in part depends on age. The color of the epidermis
is also somewhat variable, being lighter or darker green, but specimens from the
same locality are generally very uniform.

Localities in Pennsylvania represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, Horseshoe Pond, and beach-pools on Presque Isle, Erie, Erie Co.

Leboeuf Creek (outlet of lake), Waterford, Erie Co.

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MEMOIRS OF THE CARNEGIE MUSEUM.

Conneauttee Lake, Edinboro, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Lake Erie, La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).

Miami and Erie Canal, Lucas Co., Ohio (C. Goodrich).

Ohio-drainage:

Tuscarawas River, Ohio (Holland collection).

Scioto River, Columbus, Franklin Co., Ohio (H. H. Smith).

Wabash River, Bluffton, Wells Co., Indiana (C. Goodrich).

Tennessee-dramage :

Paint Rock River, Paint Rock and Princeton, Jackson Co., Alabama (H. H. Smith).

West of Mississippi:

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

Lawrence,, Douglas Co., Kansas (R. L. Moodie).

Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).

Big Deceiper Creek, Gum Springs, Clark Co., Arkansas (H. E. Wheeler).

Black Bayou, Victoria Co., Texas (D. A. Atkinson).

.1 labama-drainage :

Black Warrior River, Lock 11, Tuscaloosa, Tuscaloosa Co., Alabama (H. H. Smith).

Pond at Holt, Tuscaloosa Co., Alabama (H. H. Smith).

Beaver Creek and Canoe Creek, St. Clair Co., Alabama (H. H. Smith) (Coosa-drainage).

Othkalooga Creek, Calhoun, Gordon Co., Georgia (H. H. Smith) (Coosa-drainage).

A tlantic-dramage :

*

Greenfired Mill, Wilmington, New Hanover Co., North Carolina (G. H. Clapp donor).

Distribution and Ecology in Pennsylvania (See fig. 14) : This species is restricted
to the northwestern section of our state (Erie Co.), and is principally found in
Lake Erie, where it inhabits the sandy shores of Presque Isle Bay and the beach-
pools at the eastern end of Presque Isle. At two places I discovered it in the
Ohio-drainage, in the headwaters of French Creek, in Conneauttee Lake, in the
black muck of this lake; in the other case, at the outlet of Leboeuf Lake, on partly
muddy bottom, in slack water, and partly on gravelly bottom and on shell marl
in more strongly flowing water.

Baker (1898a) records it from muddy bottoms in creeks, small rivers, and
ponds, and Scammon (1906) calls it a lover of quiet water and muddy or somewhat
sandy banks. This agrees well with my observations.

The peculiar distribution in Pennsylvania resembles somewhat that of certain
other species, which are found onl}'' in the northwestern part of our state.

General distribution: Type locality, Ohio River (Rafinesque) .

Simpson (1900) says: “Entire Mississippi-drainage area; southern Michigan;
North Carolina to Georgia; southwest to Matamoras, Mexico.” This indicates a

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165

very wide range, but it should be qualified somewhat. It is actually found over
large parts of the Mississippi-drainage, but records are missing from the tributaries
South of the Ohio. In the Cumberland-drainage, it is only found in ponds near
Clarksville, Montgomery Co., Tennessee (Wilson & Clark, 1914); and in the
Tennessee-drainage it goes up to northern Alabama, but seems to be missing in
East Tennessee. Northward it passes into the lake-drainage in Wisconsin (Mil-
waukee, Lapham, 1860), Illinois (Baker, 1906), Michigan (Walker, 1898), Ohio
(Sterki, 1907a; Walker, 1913), and advances northeastwards to northwestern
Pennsylvania and western New York, reaching, by way of the Erie Canal, Herkimer
Co., New York (Lewis, 1860; Marshall, 1895). In the upper Mississippi, it
ascends to southeastern Minnesota (Grant, 1886; Holzinger, 1888).

Westward it occurs in the tributaries of the Mississippi in Kansas (Scammon,
1906), Arkansas, Oklahoma, and northwestern Louisiana (Vaughan, 1893; Frier-
son, 1899). It is frequent in the streams flowing into the Gulf in Texas (Singley,
1893), reaching Matamoras in Mexico, according to Simpson. In addition it is
known from the Alabama-drainage, going thence into Georgia and the Carolinas.
This section of the range is represented in the Carnegie Aluseum by specimens
from Alabama, Georgia, and North Carolina, and is probably connected westward
with the localities in Louisiana.

Genus Anodontoides Simpson (1898).

Ortmann, 1912, p. 293; Simpson, 1914, p. 466.

Type Anodonta ferussaciana Lea.

Only one species is known, which, however, has several varieties, one of which
is found in Pennsylvania.

Key to the Forms of Anodontoides.

Ui. Shell compressed, and less elongated A. ferussacianus.

a 2. Shell more swollen, more elongated, thus approaching the subcylindrical shape.

A. ferussacianus buchanensis.

Anodontoides ferussacianus (Lea) (1834).

Anodontoides ferussacianus (Lea) Simpson, 1914, p. 467.

Plate XI, fig. 5.

Records from Pennsylvania :

Ortmann, 19096, p. 195.

Characters of the shell: Shell small to medium in size, thin. Outline sub-
elliptical or subovate, moderately elongate, rounded anteriorly as well as posteriorly.

166

MEMOIRS OF THE CARNEGIE MUSEUM.

more rarely somewhat narrowed behind. Lower margin gently and rather regu-
larly curved, often nearly straight in the middle. Upper margin gently curved,
and curved down into the posterior margin, forming, or not forming a slight angle,
without a distinct wing. Beaks convex, but not much swollen, only slightly
elevated above the hinge-line, placed more or less anterior to the middle. Beak-
sculpture consisting of three to four subconcentric, fine, and rather sharp ridges,
the first two or three placed obliquely to the direction of the hinge-line (bent up
behind), the following ones showing a more or less distinct angle. On the posterior
slope there are several fine radiating lines, indicating additional bars, obliterated
on the disk. Double-looped structure of the bars is entirely absent (See beak-
sculpture of var. buchanensis, as figured by Marshall, 1890, figs. 13, 14).

Shell more or less compressed to moderately swollen, greatest convexity upon
the posterior ridge, which is rounded and not sharply marked. Posterior slope
slightly compressed.

Epidermis light green to olive, grayish, brownish or blackish, generally grayish,
yellowish, or light brown towards the beaks, mostly with numerous narrower or
wider, dark green rays, which, however, are often indistinct, and disappear in old
shells on account of the general darkening of the epidermis. Upon the posterior
slojie, there may, or may not, be a few darker, broader rays. In addition, the
epidermis is often concentrically banded, the growth rests being marked by blackish
color.

Hinge practically edentulous, straight or slightly downward curved anteriorly.
In front of the beaks it is more or less (often very indistinctly) incurved, and here
shows rudiments of cardinals in the shape of small tubercles, swellings, or mere
irregularities. Beak-cavity shallow. Dorsal muscle-scars faint, in front part of
beak-cavity. Adductor muscle-scars faint.

Nacre bluish white, sometimes discolored by greenish or creamy tints toward
the beak-cavity; more or less iridescent, chiefly 'towards the posterior end.

Shell of male and female hardly different. Although Baker (1898a, p. 73)

%

says that the shell of the female is more inflated posteriorly {i.e., at and in front
of posterior ridge) this inflation is not always present. If it is distinctly developed,
we may safely assume that we have to deal with a female, but many females do
not possess it.

L.

H.

D.

Size: 1.

Greenville, Cat. No. 61.3294

73 mm.

38 mm.

28 mm.

2.

do. “ “ do

68 “

38 “

19 “

3.

Shenango, Cat. No. 61.4207 ( 9 )

71 “

38 “

25 “

4.

do. “ “ do

62 “

35 “

18 “

ORTMANN: monograph of the naiades of PENNSYLVANIA.

167

This species grows somewhat larger in some western streams.

Soft parts (See Ortmann, 1912, p. 294). Glochidia figured by Lea (Olis. VI,
1858, PL 5, fig. 35) (immature), and. Ortmann (19116, PI. 89, fig. 12).

Breeding-season: Records for gravid females are at hand for: August 7, 1908;
Aug. 8, 1908; Aug. 26, 1911; Sept. 13, 1917; Sept. 27, 1909; Oct. 10, 1907; and
May, 1908. Glochidia have been found in September, October, and May; and
on May 14 discharging specimens have been seen.

Thus this species is bradytictic. The breeding season begins in August and
lasts till May.

Remarks: This is a species, which has a rather indifferent, generally sub-
elliptical shape, without any striking features. Externally it looks like a small
Anodonta, and the thickness of the shell is also much like that of Anodonta. More-
over the shape of the shell is rather variable, being sometimes longer, or shorter,
and more or less compressed. If the beaks are well-preserved, there is no mistake
about its identity, since the beak-sculpture is quite peculiar and characteristic.
Eroded shells may be distinguished from Anodonta by the hinge-line, which is not
so straight.

In general shape Strophitus edentulus is often much like this species, and,
when the beaks are eroded, the two species are sometimes hard to tell apart. But
they may be distinguished by the thinner shell of Anodontoides, the interior of
which is blueish white and strongly iridescent, while in Strophitus, especially in
older specimens, the nacre is more thickened toward the beak-cavity, and there
turns pure white, creamy, or even salmon-color, and is not so iridescent. Further
in Strophitus the rudiments of the cardinal teeth are more distinct, and the hinge
is more incurved in front of the beaks. When the beaks are preserved, Strophitus
is distinguished by the heavier beak-sculpture. Of course, the soft parts, chiefly
those of the female, when it is gravid, are entirely different in these two species.

The convexity of the valves varies greatly in Anodontoides ferussacianus, and
becomes sometimes rather great, thus approaching the shape of the var. buchanensis
(see below). Such specimens turn up now and then among normal ones, and
at a few localities more frequently, as for instance, in Conneaut Lake. Some
shells from this locality (and also from the outlet of the lake) might safely be called
buchanensis; while others, found with them, have the normal, more compressed
shape; similar conditions prevail in the uppermost Shenango River, at Linesville.
We actually have here the intergrades between the normal form and the variety.

It should be noted, that the form from Conneaut Creek (tributary to Lake
Erie) is the normal form, not the Lake Erie form (buchanensis).

168

MEMOIRS OF THE CARNEGIE MUSEUM.

Localities in Pennsylvania represented in the Carnegie Museum:

Shenango River, Sliarpsville, Shenango, and Jamestown, Mercer Co.; Linesville, Crawford Co.
Pymatuning Creek, Pymatuning Township, Mercer Co.

Little Shenango River, Greenville, Mercer Co.; Hadley, Mercer Co. (D. A. Atkinson).

Randolph Run, Hartstown, Crawford Co.

Conneaut Lake and Conneaut Outlet, Crawford Co.

Cussewago Creek, Mosiertown, Crawford Co. (H. &. L. Ellsworth).

French Creek, Meadville, Crawford Co.

Conneaut Creek, West Sj^ringfield, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Tributary of Lake Ontario, Cobourg, Northumberland Co., Ontario, Canada (D. Stewart).

St. Lawrence River, Clayton, Jefferson Co., New York (Miss A. H. Robin.son).^™

Genesee River, Rochester, Alonroe Co., New York (R. H. Santens).

Swan Creek, Toledo, Lucas Co., Ohio (C. Goodrich).

Cedar Creek, Jerusalem Township, Lucas Co., Ohio (C. Goodrich).

Ten Mile Creek, Silica, Lucas Co., Ohio (C. Goodrich).

Silver Creek, Williams Co., Ohio (C. Goodrich).

Otter Creek, Monroe Co., Michigan (C. Goodrich).

Raisin River, Tecumseh, Lenawee Co., Michigan (C. Goodrich).

Ohio-Mississippi-drainage:

Tuscarawas River, Ohio (Holland collection).

Winona Lake, Kosciusko Co., Indiana (E. B. Williamson).

Kishwaukee River, Rockford, Winnebago Co., Illinois (P. E. Nordgren).

Pocatalico River, Raymond City, Putnam Co., West Virginia.

Fleming Creek, Pleasant Valley, Nicholas Co., Kentucky.

Cumberland River, Orby, Bell Co., Kentucky.

Western:

Hinkston Creek, Columbia, Boone Co., Missouri (D. K. Greger).

Smoky Hill River, Logan Co., Kansas (C. AV. Gilmoi’e).

Lodgepole Creek, Julesburg, Sedgwick Co., Colorado (G. H. Clapp, donor).

Big Thompson Creek, Loveland, Larimer Co., Colorado (A. Koenig).

Distribution and Ecology in Pennsylvania (See fig. 16) : This species is found
exclusively in the northwestern section of our territory, mainly in the headwaters
of the Beaver system, and of French Creek. In addition it occurs in Conneaut
Creek, a tributary of Lake Erie. Thus it obviously falls into the same category
as some other species before mentioned. Apparently its metropolis in our state is
the upper Shenango, where it is abundant, for instance at Linesville and in Pyma-
tuning Creek. It has never been found below Sharpsville, just below the mouth
of Pymatuning Creek. In Frenc'h Creek proper it is rare (only one found at

Dwarf, pale race, possibly the var. modestus (Lea).

ORTMANN: monograph of the naiades of PENNSYLVANIA.

169

Meadville), while it is rather frequent in Conneaut Outlet and in the lake itself.
This peculiar feature will be discussed elsewhere. It suffices to mention here that^

• Anodontoides ferrussacianus.

+ Do. feruss. buchanensis.

all these systems (Beaver, Conneaut, French Creek) have been either naturally or
artificially closely connected in the past.

Wherever I found this species in abundance it preferred comparatively quiet
parts of the creeks, with little or no current, and with sandy-gravelly bottoms,
with some admixture of mud. It also occurs in lakes, as for instance Conneaut
Lake, and exhibits here the tendency to become more inflated, approaching thus
the var. buchanensis. In Conneaut Lake it is found chiefly on sandy bottom at
the south end of the lake, but also elsewhere in gravel. It seems to be averse to
larger streams with strong currents and rough bottoms.

Baker (1898a) says that it is a species frequenting muddy bottoms.

General distribution: Type locality , Ohio River, Cincinnati, Ohio (Lea).

According to Simpson this species is found generally in the Mississippi-drainage
area; in the St. Lawrence system; the Red River of the North; the Saskatchewan
River; and doubtfully in Connecticut (according to Linsley, 1845, at Whitney-
ville, New Haven Co.).

In the east its range certainly extends into 'New York state, as far as the
Hudson River and the headwaters of the Susquehanna and Delaware systems
(Marshall, 1895),^^^ and it is frequent in the St. Lawrence-drainage in a form
which is typical (see our specimens from the Genesee River). I have it also in
rather typical development from a tributary on the Canadian side of Lake Ontario,
and it has been reported from Montreal and Ottawa. Farther west, its northern
boundary is uncertain, but it surely advances rather far northward, as is shown
by the localities given by Walker (1898) for Michigan (Charlevoix, upper Penin-

But it is not found in Pennsylvania in these rivers.

170

MEMOIRS OF THE CARNEGIE MUSEUM.

siila), by localities in Wisconsin and Minnesota, and the Canadian range given by
Simpson. Southward it is well distributed over the states of Ohio to the Ohio
River (Sterki, 1907a), Indiana (Call, 1896a and 1900), and Illinois (Baker, 1906;
Forbes & Richardson, 1913). It has not been found in the Ohio above Cincinnati,
and south of the Ohio. It does not occur in southwestern Pennsylvania and in
the headwaters of the Monongahela and in the Little Kanawha in West Virginia.
But it turns up in the Big Kanawha-drainage, and is found in the Licking system
in Kentucky. It is known from the Cumberland (Wilson & Clark, 1914) (also
Carnegie Museum), and is very rare in the uppermost Tennessee system (Powell
River). (This will be discussed elsewhere.)

Westward A. ferussacianus goes across the Mississippi, where it is found in
South Dakota, Iowa, and Kansas, advancing here, in the Kansas River-drainage,
farther westward than any other shell (Decatur Co., Scammon, 1906, and Logan
Co., Carnegie Museum). In addition, the Carnegie jMuseum has received material
representing this species from the state of Colorado (Platte River-drainage), which
extends its western range here to the foot of the Rocky Mountains.

Farther south this species has not been reported, and in general the range is
rather northern.

Anodontoides ferussacianus buchanensis (Lea) (1838).
Anodontoides ferussacianus suhcylindraceus (Lea) Simpson, 1900, p. 660; Ano-
dontoides ferussacianus buchanensis (Lea) Simpson, 1914, p. 469.

Plate XI, fig. 6.

Records from Pennsylvania :

Ortinann, 19096, p. 202.^^^

Characters of variety: This form differs from the typical ferussacianus by the
more convex valves and the more elongated outline, which renders the whole shell

more nearly subcylindrical.

L.

H.

D.

Size: 1.

Presque Isle, Cat. No. 01.1577

82 mm.

39 mm.

33 mm.

2.

do. Cat. No. 61.3295

81 “

38 “

28 “

3.

do. “ “ do

75 “

38 “

29 “

4.

do. “ “ do

66 “

31 “

26 “

5.

do. “ “ do

65 “

31 “

24 “

Soft parts not different from those of the typical form. Glochidia not observed
by myself. Surber (1912, PI. 3, fig. 43) figures them; his measurements are:

Pilsbry (1894, p. 30) records “ Anodonta subcylindrica Lea ” (sic!) from York Furnace, York Co.
The specimens thus labeled in the Philadelphia Academy are Anodonta cataracta Say. (See above.)

ORTMANN: monograph of the naiades of PENNSYLVANIA.

171

0.33 X 0.33, while my measurements for the typical form (1912, p. 294) are: 0.32
X 0.32 mm.

Breeding season: Not observed. According to Surber gravid specimens have
been found in September and October.

Remarks: The subcylindrical, more elongated shape of the shell is the only
reliable character by which this form may be recognized; and even this is not
always equally well-developed. Lake Erie specimens may incline toward the nor-
mal form, and, on the other hand, more swollen individuals may turn up in the
Ohio-drainage, chiefly in the lake environment.

Special attention should be directed to the fact that in Conneaut Creek,
tributary to Lake Erie, the typical form is found; and parallel to this is the fact
that likewise from tributaries of the lake in northwestern Ohio and southeastern
Michigan this form is at hand.

In Lake Erie, principally in the beach-pools and ponds of Presque Isle, the
var. huchanensis is also characterized by its color. The latter is not greenish?
but is reddish brown, generally beautifully reddish chestnut near the beaks, shading
to dull dark brown and blackish upon the disk and toward the margins. In most
cases rays are very obscure or entirely absent, with the exception of a few indistinct
blackish ones upon the posterior slope. This color, however, certainly is only a
local, environmental effect. It resembles much the color observed in the beach-
pool form of Anodonta grandis footiana, but is more intense, and, moreover, not
all specimens from Presque Isle have it. From the bay I have specimens, which
are distinctly green, with distinct rays, and hardly any rusty brown. Such speci-
mens agree very closely with specimens from Conneaut Lake, and the connection
of the variety with the type is thus complete.

Localities represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, and beach pools of Presque Isle, Erie, Erie Co., Pennsylvania.

Pond at mouth of Elk Creek, Miles Grove (North Girard), Erie Co., Pennsylvania.

Lake Erie, Port Dover, Norfolk Co., Ontario, Canada (C. Goodrich).

Lake Erie, Buffalo, Erie Co., New York (Hartman collection).

Raisin River, Adrian, Lenawee Co., Michigan (C. Goodrich).

Lake Huron, Port Huron, St. Clair Co., Michigan (N. M. Grier).

Bessey Creek, Cheboygan Co., Michigan (H. B. Baker).

Scioto River, Kenton, Hardin Co., Ohio (C. Goodrich).

Distribution and Ecology (See fig. 16) : Type locality, Buck Creek, Ohio (Lea).^^^

In Pennsylvania this form is restricted to Lake Erie, and is found there in
great numbers, chiefly in certain beach-pools on Presque Isle. It lives there in

Location unknown". Possibly near Cincinnati?

172

MEMOIRS OF THE CARNEGIE MUSEUM.

the characteristic fine sand of the peninsula, sometimes mixed with more or less
mud.

This variety has been reported from other localities in Lake Erie, for instance
at Buffalo (Marshall, 1895), and also from Lake Michigan in Michigan (Walker,
1898) and Illinois (Baker, 1898a; Walker, 1913, p. 22). Records from other
larger or smaller lakes and from canals are abundant from eastern Canada, Lower
St. Lawrence region (according to Bell, 1859), New York (crossing over by way of
Erie Canal to the Hudson at Albany) (Marshall, 1895), northern Ohio (Sterki,
1907a), Michigan (Walker, and Baker, 1914), as far north as Schoolcraft and
Alger Cos., near Lake Superior (Winslow, 1917). But it also has been reported
from creeks and rivers, and its type-locality is a creek in Ohio. It is found in
New York, Ohio, Indiana (Call, 1896a, and 1900), and in Illinois (Baker, 1898a,
and 1906) in streams running to the lakes as well as to the Ohio. But on the
whole its range is more northeastern, and centers mainly in the drainage of the
Great Lakes and the St. Lawrence. Southward and westward it does not go
beyond Ohio and Indiana, nor beyond the northeastern corner of Illinois.

Genus Alasmidonta Say (1818).

Ortmann, 1912, p. 294; Simpson, 1914, p. 492; Ortmann, 1914, p. 44.

Tyjie Monodonta undulata Say.

This genus has been divided into subgenera by Simpson, but they have been
somewhat modified by the present writer. Three subgenera are found in Penn-
sylvania.

Key to the Subgenera.

ai. Hinge complete, with pseudocardinals and laterals, but the latter reversed, two in right, one in left

valve. Beak-sculpture moderately heavy Subgenus Prolasmidonta.

a 2. Hinge incomplete, laterals obliterated, pseudocardinals present. Beak-sculpture very heavy.

bi. Pseudocardinals well-developed, stumpy. Posterior ridge weak, posterior slope smooth (rarely

with traces of wrinkles). Rays not broken up into spots Subgenus Alasmidonta.

62. Pseudocardinals weak, compressed. Posterior ridge distinct, posterior slope more or less cor-
rugated. Rays generally broken up into spots Subgenus Decurambis.

Subgenus Prolasmidonta Ortmann (1914).

Ortmann, 1914, p. 44.

Type Unto heterodon Lea.

There is only one species in this sub genus.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

173

Alasmidonta (Prolasmidonta) heterodon (Lea) (1830).
Alasmidonta heterodon (Lea) Simpson, 1914, p. 499; Alasmidonta {Prolas7nidonta)
heterodon (Lea) Ortmann, 1914, p. 44.

Plate XII, figs. 1, 2.

Records from Pennsylvania:

Lea, 1830 (Schuylkill River and Darby Creek, the latter chiefly in Delaware Co.)

Conrad, 1838 (Schuylkill River).

Gabb, 1861 (Schuylkill River, below Fairmont Dam, Philadelphia.)

Hartman & Michener, 1874 (Schuylkill River, Chester Co.)

Marshall, 1895 (Philadelphia).

Schick, 1895 (Schuylkill Canal at Philadelphia and Manayunk; Neshaminy Creek, Bucks Co.)

Ortmann, 19096, p. 207. *

Characters of the shell: Shell small and rather thin. Outline subrhomboidal
or subtrapezoidal to subovate, and more or less elongate, rounded anteriorly,
subangular behind, with an upper posterior angle (which may be obsolete), and a
more distinct and produced lower posterior angle. Lower margin gently curved
(chiefly in the male), or straight and even slightly concave (chiefly in the female).
Beaks somewhat inflated, but not very prominent, anterior to the middle of the
shell, but not very near the anterior end. Beak-sculpture consisting of three or
four (rarely traces of a fifth) bars, the first two concentric and simple, the following
ones with a distinct angle upon the posterior ridge, in front of which is a more or
less distinct, shallow sinus. The bars are moderately heavy and blunt, and the
sinus never assumes the shape of a re-entering angle, sometimes it is barely indi-
cated. Thus the beak-sculpture cannot be called double-looped. Upon the
posterior slope there are one or two fine radiating lines, indicating as many addi-
tional bars, which are obliterated on the disk.

Shell more or less swollen and inflated, chiefly in the region of the posterior
ridge, and in the female; the sides are rather flat. Posterior ridge quite distinct,
generally more so in the female than in the male, but also in the latter this ridge
is distinct, although rounded. In the female, it becomes almost angular. The
posterior slope is somewhat compressed in the male, while in the female it appears
sometimes almost truncated.

Epidermis greenish olive to brownish. Sometimes obscure (rarely more dis-
tinct), simple and straight rays are present, but in other cases, rays are invisible,
and the color of the whole surface is uniform. Often there are indistinct con-
centric bands of lighter and darker green. Growth-rests indistinct.

Hinge complete, with pseudocardinals and laterals, but teeth rather feeble.
Pseudocardinals two in each valve, compressed and crenulated, often the anterior

174

MEMOIRS OF THE CARNEGIE MUSEUM.

in the right valve, and sometimes the posterior in the left valve rudimentary.

In addition, there is an interdental tooth in the left valve, which may be well-
developed and isolated, or may be more or less connected with the posterior pseudo-
cardinal, or may be even rudimentary. There is great variation in this respect.
Laterals rather thin and short; characteristically there are two laterals in the
right, and one in the left valve (just the reverse of the normal condition in the
Naiades), but there is considerable variation in the laterals. The upper one is
generally not so sharp as the lower one, and the former may be more or less rudi-
mentary, sometimes only for a part of its length. Beak-cavities moderately
developed, not deep. Dorsal muscle-scars distinct, in the beak-cavity. Adductor
muscle-scars moderately deep anteriorly, faint posteriorly.

Nacre bluish or silvery white, iridescent, and often inclining toward cream-
color or yellowish in the beak-cavity; sometimes with greenish or grayish dis-
coloration.

Shells of males and females distinguishable. The male as a rule has a more
compressed, and more ovate and elongate shell, with the lower margin gently and
uniformly curved, the posterior ridge less sharp, and the posterior slope not trun-
cate; while in the female the shell is more swollen in the region of the posterior
ridge and just in front of it; has a more distinct posterior ridge, which renders
the posterior slope more truncate, and makes the whole outline of the shell shorter
and more trapezoidal. Furthermore in the female the lower iiosterior angle is .
more produced downward, so that the lower margin of the shell becomes rather
straight, in some cases even concave. In young specimens the sexual differences
of the shell are not seen, and prevalently the young females in shape resemble males.

L. H. D.

Size: 1. Kunkletown, Cat. No. G1.4652 (cT) 48 mm. 27 mm. 19 mm.

2. Manayunk, Cat. No. 61.4250 (d") 42 “ 23 “ 14 “

3. do. Cat. No. 61.4249 (9 gravid) 40 “ 23 “ 16 “

4. do. Cat. No. 61.4250 (o') 38 “ 21 “ 14 “

5. do. “ “ do. (9 gravid) 38 “ 21 “ 15 “

6. do. Cat. No. 61.4249 ( 9 gravid) 35 “ 20 “ 14 “

tiojt parts (See Ortmami, 1912, p. 295). Glochidia (See Ortmann, 19116, PI.
89, fig. 8).

Breeding season: Cornier (1909, p. 112) found this species gravid in February,
and I found gravid females with glochidia on April 24, 1909.

Remarks: A small species, easily overlooked, with no striking external features.
However, the general shape, chiefly the distinct posterior ridge, give it the un-

ORTMANN: monograph of the naiades of PENNSYLVANIA.

175

mistakable aspect of an Alasmidonta. If attention is paid to the hinge, there can
be no mistaking this shell, for the teeth are unique, at least among Pennsylvanian
shells.

There are no species related to this in western Pennsylvania, and this species
has altogether a rather isolated position in the system (See Ortmann, 1913a, pp.
324, 361).

Localities represented in the Carnegie Museum:

Delaware River, Shawnee, Monroe Co., Pennsylvania.

Princess Creek, Kiinkletown, Monroe Co., Pennsylvania.

Schuylkill Canal, Manayunk, Philadelphia Co., Pennsylvania.

Marsh Run, three miles S. E. of Remington, Fauquier Co., Virginia (Rappahannock-drainage).
Mountain Run, Culpeper, Culpeper Co., Virginia (Rappahannock-drainage).

Locality represented in the Philadelphia Academy of Natural Sciences:

Big Neshaminy Creek, “ Edderton,” Bucks Co., Pennsylvania (H. W. Fowler).

Distribution and Ecology (See fig. 17) : Type locality, Schuylkill River, Penn-
sylvania (Lea) .

In Pennsylvania this species is decidedly rare. I have found it in large numbers
only in the Schuylkill Canal; at other localities only a few were obtained. It is

Alasmidonta heterodon.

• Alasmidonta undulata.

known from a number of localities in the Delaware-drainage in the vicinity of
Philadelphia, and chiefly from streams on the Piedmont Plateau. But in the

Probably = Eddington.

176

MEMOIRS OF THE CARNEGIE MUSEUM.

Delaware at Shawnee and at Kimkletown (in a stream tributary to the Lehigh
River) I found it to the northwest of the Blue Mountain (Kittatinny Mountain).

No trace of it has ever been seen in the Susquehanna and Potomac drainages,
but it is positively present in the Rappahannock-drainage in Virginia, and here
again it is on the Piedmont Plateau. (Ortmann, 1913a, p. 319.)

According to Simpson (1900), it goes from ‘biorthern New England to Vir-
ginia,” but even in New England and New York locality-records are quite scarce.
It is known from the Connecticut River, Hartland, Windsor Co., Vermont (Mar-
shall, 1895), from Massachusetts (Marshall, but not mentioned by Gould-Binney,
1870), and in Connecticut from the Housatonic River (Linsley, 1845) and Mixville
(Cheshire) New Haven Co. (Marshall) (see also Johnson, 1915). In New York,
it is known from the southeastern portion (IMarshall, without exact localities).
Otherwise definite records are missing, but the extension of its range southward to
Virginia, as given by Simpson, has been confirmed by the present writer with regard
to the Rappahannock River.

For the present it may be said that the chief feature of the distribution of
this species is its erratic character. As far as I can now see, it seems to be most
likely a species of the Piedmont Plateau. Its apparent absence in the Susquehanna
and Potomac drainages may be due to the incomjilete investigation of the Piedmont
tributaries of these rivers.

At Kimkletown and in the Rappahannock-drainage, I found this species in
very small streams (‘'runs”), in strongly flowing water, and in rather coarse
gravel, but at each locality only few specimens. In the Delaware at Shawnee it
was (one specimen) in a small branch of the river in sand and moderate current.
In the only case where I collected large numbers (aliout fifty), I found it under
very different conditions, and that was in the canal at Manayunk at a time when
the water had been drained off and only a few inches of water were left in the
middle. Here the bottom consisted of larger and smaller stones, the interstices
hlled with sandy mud. This, however, cannot be regarded as normal, and the
ecological conditions of this species remain to be studied.

'Subgenus Alasmidonta Simpson (1900).

Almmidonta Simpson, 1914, p. 493, + Bullella, ibid., p. 508; Alasmidonta Ort-
mann, 1914, p. 45.

Type Alonodonta undulata Say.

Only one species in Pennsylvania belongs to this subgenus.

ORTMANN: monograph op the naiades of PENNSYLVANIA.

177

Alasmidonta (Alasmidonta) undulata (Say) (1817).^^^

Alasmidonta undulata (Say) Simpson, 1914, p. 494.

Plate XI, fig. 7.

Records from Pennsylvania:

Say, 1817 (Delaware River).

Gabb, 1861 (Delaware, Schuylkill, and Wissahickon, Philadelphia; headwaters of Frankford Creek,
Montgomery Co.)

Bruckhart, 1869 (Lancaster Co.)

Hartman & Michener, 1874 (Chester Co.)

Harn, 1891 (western Pennsylvania).^^®

Schick, 1895 (Canal at Manayunk, Philadelphia Co.; Munckinipattus Creek, Glenolden, Delaware Co.;

Neshaminy Creek, Bucks Co.)

Ortmann, 19096, p. 207.

Characters of the shell: Shell rather small, and rather thick, chiefly in its
anterior part. Outline subtriangular, siibovate, or almost subelliptical; generally
comparatively short and high, more or less narrowed and pointed behind. Lower
margin more or less convex. Upper margin straight, forming or not forming an
angle with the obliquely descending posterior margin. Beaks more or less inflated,
and somewhat elevated above the hinge-line, situated more or less anteriorly.
Beak-sculpture extremely heavy, but somewhat variable. There are specimens with
much heavier sculpture than others. Sculpture consisting of four or five bars,
which are angular and much elevated posteriorly, straight or very little sinuated
in the middle, rounded and low anteriorly, running up to the beaks upon the
posterior slope as fine, but distinct, converging lines, and, in addition, there are
several similar radiating lines behind them, as well as in front of the beaks. (See
Marshall, 1890, fig. 11.) Upon the posterior slope there are sometimes irregular,
oblique corrugations, but in most cases, they are absent.

Shell more or less swollen, lateral faces gently convex, with an oblique pos-
terior ridge, which is rounded and often indistinct. Posterior slope somewhat
compressed, rarely subtruncate.

Epidermis greenish, yellowish, reddish-brown, brown, or black, with more or
less distinct dark green to blackish rays, which are broader or narrower, straight,
uninterrupted. Old shells are often uniformly black-brown. Often the posterior
slope is lighter than the rest of the shell, with more distinct, sharper, and finer
rays. Frequently there are lighter or darker concentric bands. Growth-rests
rather indistinct.

125 Not 1816.

12® This species certainly is absent from the Ohio-drainage. Ham’s specimens may have come from
the headwaters of the West Branch of the Susquehanna (See Ortmann, 19096, p. 180).

178

MEMOIRS OF THE CARNEGIE MUSEUM.

Hinge incomplete. Pseudocardinals present, stumpy, crenulated, generally
one in right, two in left valve, of which the anterior may be rudimentary. Inter-
dental tooth of left valve poorly developed, generally connected with the posterior
pseudocardinal. Laterals practically absent. Beak-cavity moderately deep.
Dorsal muscle-scars distinct, upon the hinge-plate. Adductor scars deeply im-
pressed anteriorly, less so posteriorly. Nacre white, salmon, pink, or red, the red
shades prevailing, very iridescent posteriorly.

Shell of male and female indistinguishable.

L. H. D.

Size: I. Kimberton Dam, Cat. No. 61.883 77 mm. 52 mm. 37 mm.

2. Tioga, Cat. No. 61.4242 (9 gravid) 69 “ 41 “ 28 “

3. do. “ “ do. (d") i 66 “ 34 “ 23 “

4. Greeiicastle, Cat. No. 61.4247 (9 gravid) 48 “ 32 “ 18 “

5. do. “ “ do. ((y) 42 “ 27 “ 16 “

Soft parts (See Ortmann, 1912, p. 296). Glochidia (See Ortmann, 19116,
PI. 89, fig. 9).

Breeding season: I have the following dates for gravid females: July 18,
1908; July 22, 1910; Aug. 10, 1910; Aug. 11, 1910; Aug. 12, 1908; Aug. 13,
1910; Aug. 19, 1909; Sept. 5, 1909; Sept. 6, 1909. Then in spring, April 24,
1909; April 26, 1909; May 6, 1912; May 10, 1912; May 11, 1912; June 7, ;912;
June 12, 1912; June 14, 1910.

This is clearly a hradytictic form, with a rather short interim, in June and July.
Remarks: Quite a well characterized species, not to be confounded with any
other Pennsylvanian form, and having no related forms in the interior drainage
system. The most nearly allied species are found in the southern parts of the
Atlantic watershed. South Carolina and Georgia. It is easily recognized by the
shape of the shell, heavy beak-sculpture, by the color and markings of its epi-
dermis, and, on the inside of the shell, by the peculiar character of the hinge,
and the often beautifully red tints of the nacre.

It is a rather variable shell, and the chief variations have been alluded to
above.

Pennsylvanian localities, represented in the Carnegie Museum:

Delaware-drainage :

Delaware River, Yardley, Bucks Co.; Shawnee, Monroe Co.

White Clay Creek, Avondale, Chester Co.

Schuylkill Canal, Manayunk, Philadelpliia Co.

Kimberton Dam, Chester Co. (Hartman collection) (formed by French Creek, near Phoenixville, Chester
Co.)

Princess Creek, Kunkletown, Monroe Co.

ORTMANN: monograph op the naiades op PENNSYLVANIA.

179

Susquehanna-drainage :

Susquehanna River, Selinsgrove, Snyder Co.

Muddycreek, Lancaster Co. (G. H. Clapp, donor) (drainage of Conestoga Creek).

Conewago Creek, Table Rock, Adams Co.

Conodoguinet Creek, Carlisle, Cumberland Co.

Frankstown Branch Juniata River, Hollidaysburg, Blair Co.

Raystown Branch Juniata River, Everett, Bedford Co.

Shobers Run, Bedford Springs, Bedford Co. (A. Koenig).

West Branch Mahantango Creek, Richfield, Juniata Co. (D. A. Atkinson).

Beaver Dam Creek, Flinton, Cambria Co. (D. A. Atkinson).

Swartz Run, Ashville, Cambria Co.

Chest Creek, Patton, Cambria Co.

North Branch Susquehanna River, Tuhkhannock, Wyoming Co.

Chemung River, South Waverly, Bradford Co.

Mill-race of Crooked Creek, Tioga, Tioga Co.

Potomac-drainage :

East Branch Little Antietam Creek, Waynesboro, Franklin Co.

Conococheague Creek, Greencastle and Scotland, Franklin Co.

West Branch Conococheague Creek, Mercersburg Junction, Franklin Co.

Great Tonoloway Creek, Thompson Township, Fulton Co.

Pennsylvanian localities represented in the Philadelphia Acadeiny of Natural
Sciences:

Manatawny Creek, Earlville, Berks Co. (H. A. Pilsbry).

Sacony Creek, Kutztown, Berks Co. (H. K. Deisher).

Schuylkill River, Phoenixville, Chester Co. (C. M. Wheatley).

“Valley Creek,” southwest of Coatesville, Chester Co. (C. H. Conner).

“ Big Elk Creek, Westgrove,” Chester Co. (C. H. Conner).

Lancaster, Lancaster Co. (J. B. Eshleman).

Pequea Creek, Paradise, Lancaster Co. (W. Stone).

Susquehanna River, York Furnace, York Co. (W. Stone).

I

Other localities, represented in the Carnegie Museum:

Aroostook River, Caribou, Aroostook Co., Maine (0. 0. Nylander).

Fish River, Eagle Lake, Aroostook Co., Maine (0. 0. Nylander).

Delaware River, Newbold, Gloucester Co., New Jersey (C. H. Conner).

Potomac River, Hancock, Washington Co., Maryland.

Wills Creek, Ellerslie, Allegany Co., Maryland.

South Branch Potomac River, Romney, Hampshire Co., West Virginia.

Shenandoah River, Harpers Ferry, Jefferson Co., West Virginia.

North Fork Shenandoah River, Broadway, Rockingham Co., Virginia.

South Fork Shenandoah River, Elkton, Rockingham Co., Virginia.

Probably Sucker Run, tributary to West Branch Brandywine Creek.

Not exact. The Middle Branch of White Clay Creek is at Westgrove and Big Elk Creek is
about four or five miles farther West, at Lincoln University.

180

MEMOIRS OP THE CARNEGIE MUSEUM.

South River, Waynesboro, Augusta Co., Virginia.

Rappahannock River and Marsh Run, Remington, Fauquier Co., Virginia.

North River, Buena Vista and Le.xington, Rockbridge Co., Virginia.

Calf Pasture River, Goshen, Rockbridge Co., Virginia.

Distribution and Ecology in Pennsylvania (See fig. 17) : A common species in
the Atlantic-drainage in Pennsylvania; but it is quite evident that it avoids the
larger rivers, and prefers the smaller streams, where it becomes locally very abun-
dant, going far up towards the headwaters, as is best seen in the upper Juniata and
the tributaries of the West Branch of the Susquehanna in Cambria Co. Here it
closely approaches the divide, but in no instance has it been found west of the
divide.

It does not seem to favor riffles and very rough water, but is found chiefly in
more (piiet iiarts, but with some current, for instance, above riffles, where a steady
flow of water prevails. It does not like slackwater, but occasionally it is found
in ponds and canals; and it has been reported from several lakes in New York.
It also likes mill-races, if the current is not too rapid. It lives mostly in a mixture
of coarser or finer gravel with sand and mud; but I have taken it also in eddies
with slow current embedded in the mud deposited between larger stones.

General distribution: Type locality, Delaware River, near Philadelphia (Say).

According to Simpson (1900), this species extends from the Lower St. Law-
rence-drainage southward to North Carolina.” Its presence in the lower St.
Lawrence (below Lake Ontario) is well established (Bell, 1859; Whiteaves, 1863;
Marshall, 1895). In New York state it is found in the St. Lawrence-drainage,
as in Lake Champlain (De Kay, 1843), Seneca and Crooked Lakes (Dewey, 1856);
and in Onondago Co. (Marshall). It is more abundant still in the drainage of
the Mohawk and Hudson Rivers, and of the upper Susquehanna (Chenango,
Canisteo, Conhocton, and Tioga Rivers) (See De Kay, 1843; Lewis, 1860; Marshall,
1895).

It has been reported from Maine (Lermond, 1909; Nylander, 1914), from
Keene, Cheshire Co., New Hampshire (Walker & Coolidge, 1908), and is rather
common in Vermont (Adams, 1842), Massachusetts (Gould-Binney, 1870), Rhode
Island (Carpenter, 1890), Connecticut (Linsley, 1845; Perkins, 1869) (See also
Johnson, 1915, p. 26).

South of Pennsylvania it has been previously recorded from Sideling Creek,
Allegany Co., Maryland (Pilsbry, 1894), and from Raleigh, Wake Co., North
Carolina (Marshall, 1895), but other exact localities are not knowm to me. How-
ever that it exists in the Potomac, Rappahannock, and upper James drainages
is shown by the material collected by myself for the Carnegie Museum.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

181

The exact southern boundary of this species in North Carolina is unknown.
Although its metropolis seems to be in the northern parts of the Atlantic watershed,
we have reason to believe, on account of its southern affinities, that it belongs to
the southern element in the Atlantic fauna (Ortmann, 1913a, pp. 324 and 361).

Subgenus Decurambis Rafinesque (1831).

Rugifera Simpson, 1900, p. 670; 1914, p. 504; Ortmann, 1914, p. 46; Decurambis
Frierson, 1914a, p. 7.

Type Alasrnodonta marginata Say.

Two species and one variety belong here.

Key to the Forms of Decurambis.

«i. Posterior slope strongly truncate. Beaks placed more centrally, anterior end of shell more produced.
hi. Shell greenish, posterior slope slightly lighter than the rest, but generally with rays.

A. (D.) marginata.

hi. Shell with posterior slope contrasted wuth the rest, light or reddish brown, hardly with rays.
Rest of the shell brownish or yellowish, with distinct green rays.

A. (D.) margmata susquehanncB.

a 2. Posterior slope less strongly truncate. Beaks placed more anteriorly, and anterior end of shell
shorter A. (D.) varicosa.

Alasmidonta (Decurambis) marginata (Say) (1819).

A. truncata (B. H. Wright) Simpson, 1900, p. 671; A. marginata (Say) Fox,
1901, p. 47; Alasmidonta 7narginata (Say) Simpson, 1914, p. 504.

Plate XII, fig. 3.

Records from Pennsylvania:

Harn, 1891 (western Pennsylvania).

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co.; Beaver River, Wampum, Lawrence Co.)
Ortmann, 19096, p. 196.

Characters of the shell: Shell of medium size, moderately thick, but rather thin
when young. Outline subtrapezoidal, rounded before and triangular behind.
Lower margin gently convex, more or less straight, or even somewhat concave in
the posterior part. Upper margin straight or gently convex, forming a more or
less distinct angle with the obliquely descending posterior margin. Beaks large
and inflated, somewhat elevated above the hinge-line, situated somewhat in front
of the middle, but at a considerable (although variable) distance from the anterior
end. Beak-sculpture heavy, consisting of three or four thick bars, the second and
third being a little produced and angular upon the posterior ridge, and having a
more or less distinct sinus in front of the angle. This sinus is quite variable, some-

182

MEMOIRS OF THE CARNEGIE MUSEUM.

times rather distinct, so as to render the bars almost double-looped, sometimes
hardly indicated, so that the middle part of the bars is almost straight. The
fourth bar, if present at all, is rudimentary, and only its middle part is distinct.
Radiating lines upon the posterior slope indistinct or absent. But upon the
posterior slope there is a system of fine, irregular, radial wrinkles or ridges, running
toward the upper posterior margin. These wrinkles are rarely absent or obscure,
but generally well-developed; they may be finer or coarser, and may extend over a
larger or smaller portion of the posterior slope; generally they are less distinct
toward the posterior end in old shells.

Shell inflated, greatest width at or immediately in front of the posterior ridge.
Lateral faces gently convex, with a very distinct posterior ridge, which, although
rounded, is well marked in consequence of the truncated character of the posterior
slope. Very often the iiosterior ridge appears as if bounded by two ridges, includ-
ing an elevated part of the shell between them: this is chiefly the case toward the
posterior end of old shells. The posterior slope is somewhat compressed and
elevated in the middle, but in its general shape it presents a truncate, in some
cases almost flat appearance.

Epidermis yellowish, greenish, or brownish, blackish in old shells, generally
with very distinct greenish or blackish rays. Rays narrow or wide, straight,
in most cases more or less broken up into (or overlaid by) darker spots. The
spots are best developed towards the beaks, but often they are distinct all over
the shell, and very rarely entirely wanting. Generally, the posterior slope is
lighter in color than the rest of the shell, and has fewer and less distinct rays and
spots. Often there are also lighter and darker concentric bands, marking, more
or less, the growth-rests. In old shells, the color pattern becomes obscure, and
the shell is more or less uniformly blackish.

Hinge incomplete. Pseudocardinals rather thin, but distinct, depressed-
triangular (not stumpy), one in each valve, and the left valve has, in addition, a
more or less well-developed interdental tooth, which may be separated from the
pseudocardinal, or more or less united with it. More rarely it is rudimentary.
Lateral teeth absent. Beak-cavity moderate. Dorsal muscle-scars in the beak-
cavity upon the hinge-plate. Adductor-scars distinct and rather deeply impressed
anteriorly, less so posteriorly.

Nacre bluish white, sometimes in .places greenish or grayish, discolored,
iridescent posteriorly, rarely with very pale salmon tints, never distinctly pinkish
or reddish.

Shell of the female hardly distinguishable from that of the male. Some

ORTMANN: monograph of the naiades of PENNSYLVANIA.

183

shells are more inflated over the posterior ridge than others, and when this swelling
reaches considerable proportions, we may be assured that we have to deal with a
female. But not all females have this inflation, and often males are more inflated
than females of the same size.

DLstance of

beaks from

Pr.ct. of

L.

H. D.

anterior end.

length.

1. Shenango, Cat. No. 61.4270 (cf )

. .96 mm. 56

mm. 35 mm.

34

.35

2.

do. “ “ do. ( 9 gravid) . . .

..92 “ 53

“ 40 “

29

.32

3.

Rose Point, Cat. No. 61.3139 (cf)

..81 “ 42

00

CO

26

.32

4.

do. “ “ do. ( 9 gravid) . .

..73 “ 39

“ 27 “

22

.30

5.

do. “ “ do. (cf)

..68 “ 39

“ 28 “

23

.34

6.

do. “ “ do. (9 gravid)..

..63 “ 36

“ 26 “

21

.33

From the above table it is seen that the females are not always more swollen
than the males. The location of the beaks is at about one-third of the length of
the shell from the anterior end.

Soft parts (See Ortmann, 1812, p. 297). Glochidia figured by Lea (Obs. VI,
1858, PI. 5, fig. 27) and Surber (1912, PL 3, fig. 42). Surber’s measurements are:
0.35 X 0.38, while mine are: 0.33 X 0.36 mm.

Breeding season: My records for breeding females cover the time from July 19
(eggs) till October 23. The earliest date for glochidia is September 2. It is to

Fig. 18.

• Alasmidonta marginata.

+ A. marginata susquehannce.

be regretted that no dates are at hand for the spring months, but the species is
undoubtedly bradytictic.

184

MEMOIRS OF THE CARNEGIE MUSEUM.

Remarks: This species is easily recognized and distinguished from others of
the genus by the general shape, sculpture of the posterior slope, color of the epi-
dermis, and by the hinge-teeth. There is no anodontine shell, except Alasmidonta
varicosa, which could be confounded with it, but it bears an external and super-
ficial resemblance in shape and color to a lampsiline shell, Truncilla triquetra
Rafinesque. The latter, however, is much smaller and has a different hinge.

The typical form of A. rnargmata is a western shell. It is rather variable in
shape. Its chief characters are its size, strongly truncate posterior end, and,
in consequence of this, the more median position of the beaks, with the anterior
end of the shell more develojied. In this it is distinguished from its eastern repre-
sentative, A. varicosa. But there are shells found in western Pennsylvania, chiefly
in the mountain-streams, in which the truncation of the posterior end is not so
lironounced. The existence of a peculiar race in the Susquehanna-drainage will
be discussed below (See under var. susquehannce) .

Specimens from Conneaut Creek (drainage of Lake Erie) do not differ from
the tyjiical A. marginata.

Pennsylvanian localities represented in the Carnegie Museum:

Ohio- and Beaver-drainages:

Little Beaver Creek, Cannelton, Beaver Co. (Miss Vera White & IT. 11. Sinitli).

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. IL Smith).

Comioquenessing Creek, Ellwood Citjq Lawrence Co. (G. H. Clapp & H. H. Smith).

Sliirpeiyrock Creek, Wurtemberg and Rose Point, Lawrence Co.

Wolf Creek, Grove City, Mercer Co.

Mahoning River, Mahoningtown, Lawrence Co.

Neshannock Creek, Eastbrook and Volant, Lawrence Co.; Ijeesburg, IMercer Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsville, Clarksville, Shenango, and
Jamestown, Mercer Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

Little Shenango River, Greenville, Mercer Co.

A llegheny-drainage :

Allegheny River, Godfrey, Kelly, and Templeton, Armstrong Co.; Walnut Bend, Venango Co.; Tionesta,
and Hickory, Forest Co.; Warren, Warren Co.; Larabee, McKean Co. (Dennis DalljO-
Buffalo Creek, Harbison, Butler Co.

Loyalhanna River, Idlepark and Ligonier, Westmoreland Co.

Quemahoning Creek, Stanton’s Mill, Somerset Co.

Crooked Creek, Rosston, Armstrong Co.

Little Mahoning Creek, Goodville, Indiana Co.

French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co.
Leboeuf Creek, Waterford, Erie Co.

Connewango Creek Russell, Warren Co.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

185

M onongahela-drainage :

Cheat River, Cheat-Haven, Fayette Co.

Lake Erie-drainage:

Conneaut Creek, West Springfield, Erie Co.

Other localities represented in the Carnegie Museum :

Lake-drainage :

Conestogo River, Conestogo, Waterloo Co., Ontario,' Canada (Miss Maria Jentsch).'^®

Sandusky River, Upper Sandusky, Wyandot Co., Ohio (C. Goodrich).

Maumee River, Waterville, Lucas Co., and Defiance, Defiance Co., Ohio (C. Goodrich); Fort Wayne,
Allen Co., Indiana (C. Goodrich).

Ohio-drainage :

Tuscarawas River, Ohio (Holland collection).

Cheat River, Jaco and Mount Chateau, Monogalia Co., West Virginia.

West Fork River, Lynch Mines, Harrison Co.; Weston, Lewis Co., West Virginia.

Little Kanawha River, Grantsville, Calhoun Co., West Virginia (W. F. Graham).

North Fork Hughes River, Cornwallis, Ritchie Co., West Virginia.

Elk River, Shelton, Clay Co.; Gassawaj^, Braxton Co., West Virginia.

Greenbrier River, Ronceverte, Greenbrier Co., West Virginia.

Reed Creek, WjTheville, Wythe Co., Virginia.

Tennessee-drainage :

Bear Creek, Burleson, Franklin Co., Alabama (H. IT. Smith).

Shoals Creek, Lauderdale Co., Alabama (H. H. Smith).

Elk River, Fayetteville, Lincoln Co., Tennessee (H. H. Smith).

Flint River, Gurley, Madison Co., Alabama (G. H. Clapp, donor).

Paint Rock River, Paint Rock and Trenton, Jackson Co., xllabama (H. H. Smith).

Holston River, Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill and Holston Station, Grainger
Co.; Austin Mill, Hawkins Co., Tennessee.

South Fork Holston River, Pactolus, Bluff City, and Emmett, Sullivan Co., Tennessee.

Middle Fork Holston River, Chilhowie, Smyth Co., Virginia.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia; Mendota,
Washington Co., Virginia; Saltville, SmjTh Co., Virginia.

Big Mocassin Creek, Mocassin Gap, Scott Co., Virginia.

Clinch River, Edgemoor and Clinton, Anderson Co.; Black Fox Ford, Union Co.; Clinch River Station,
Claiborne Co.; Oakman, Grainger Co., Tennessee; Speers Ferry and Clinchport, Scott Co., Vir-
ginia; St. Paul, Wise Co., Virginia; Fink and Cleveland, Russel Co., Virginia; Richland, Tazewell
Co., Virginia.

Powell River, Combs, Claiborne Co., Tennessee; Dryden, Lee Co., Virginia.

West of Mississippi:

Gasconade River, Gasconade, Gasconade Co., Missouri (W. 1. Utterback).

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

White River, Cotter, Baxter Co., Arkansas (A. A. Hinkley).

Grand River-drainage, to Lake Erie. The specimen at hand is normal.

186

MEMOIRS OF THE CARNEGIE MUSEUM.

Distribution and Ecology in Pennsylvania (See fig. 18) : This species is rather
frequent in the Ohio-drainage in western Pennsylvania, but decidedly avoids the
larger rivers, and is more or less absent (probably extinct) in the Monongahela-
drainage. Rhoads mentions it from the Ohio at Coraopolis, below Pittsburgh,
but I never found it in the Ohio proper, nor in the Allegheny below Armstrong Co.
In the Ohio between Pennsylvania and Cincinnati it is also missing. In the
Monongahela-drainage it has been found only in Cheat River, but it turns up again
in the upper Alonongahela system (Cheat and West Fork Rivers) in West Virginia.

In the Beaver-drainage, this species is abundant, except in the extreme head-
waters. From Armstrong County upwards it is found regularly in almost all
streams tributary to the Allegheny, and goes far up into the upper Loyalhanna in
Westmoreland Co., the Quemahoning in Somerset Co., the Little Mahoning in
Indiana Co., and the uppermost Allegheny in McKean Co.

A. marginata is most decidedly a species of the riffles, being found there in
finer or coarse, but firmly packed gravel, in swift currents. It is one of the species,
which in the Conemaugh-drainage enter the valleys between Chestnut and Laurel
Ridges and the Allegheny Front.

It occurs also in Coimeaut Creek of the Lake Erie-drainage. Here it is
rather abundant, and does not differ at aU from the common form of western
Pennsylvania. It should be noted that it never has been found on the Pennsyl-
vanian shores of Lake Erie, although Sterki (1907a) reports a small, slight form,
from the lake in Ohio, and Walker (1913, p. 22) reports A. varicosa from Lake
Erie. I have never seen this lake-form.

Call (1900) remarks on the strongly developed foot of this species, which is,
indeed, quite remarkable. When fully extended, the foot is firmly attached to
the gravel in the river bed, and it requires quite an effort to dislodge the specimens.

General distribution: Type locality, Scioto River, Chillicothe, Ross Co., Ohio,
Say. (See Fox, 1901.)

Simpson (1900) gives this species as from the Upper Mississippi-drainage;
Ohio, Cumberland, and Tennessee river systems; Michigan; Upper St. Lawrence-
drainage.” This is about right, but it should be noted that the species advances
eastward into certain mountain-streams of the western slope of the AUeghenies,
not only in the headwaters of the Tennessee in Tennessee and Virginia, but also
in West Virginia and Pennsylvania, in the upper Kanawha system, and in the
. upper Conemaugh. In the Allegheny it also goes far up (to Olean, Cattaraugus
Co., New York) (Marshall, 1895) and to McKean Co., Pennsylvania. It is known
from several other localities in western New York, partly in the St. Lawrence,

ORTMANN: monograph of the naiades of PENNSYLVANIA.

187

partly in the Atlantic-drainage (Marshall and Baker, 18986), but. these records
should be confirmed on account of the possibility, that this form may have been
confounded with the eastern forms. Thus the northeastern boundary of A.
77iarginata is yet obscure. Northward this species has certainly crossed over into
the lake-drainage, as is shown by a number of localities represented in the Carnegie
Museum, and has been reported for Ohio (Dean, 1890; Sterki, 1907a), Indiana
(Call, 1896a and 1900), and Illinois (Baker, 1898a, 1906). In Michigan, it has
spread over the southern parts of the state, as far north as Roscommon Co. (Walker,
1898), and it is present in Ontario, north of Lake Erie (Carnegie Museum).

In the Mississippi this species goes up from Iowa (Pratt, 1876; Witter, 1878)
to Minnesota (Grant, 1886; Holzinger, 1888).

West of the Mississippi, it seems to be much rarer. It is in the Wapsipinicon
and Volga Rivers in Iowa (Geiser, 1910), and at Iowa City, Johnson Co., Iowa
(Marshall), in Missouri (Utterback, 1916), but not in Kansas (Scammon, 1906).
It is in the Ozark region of northern Arkansas (Carnegie Museum and Meek &
Clark, 1912) going to the Ouachita River in central Arkansas (Wheeler, 1918).
Thence southward it is missing.

Alasmidonta (Decurambis) marginata SUSQUEHANNA Ortmann.
Ortmann, 19136, p. 315 {et passim, sine descriptions).

Plate XII, fig. 4.

«

Not previously recorded, except by Ortmann (19I3&).

Characters of variety: Shell somewhat smaller than that of the normal form.
Color of epidermis peculiar, brighter. Epidermis more or less brown (pale or
reddish’ brown), with distinct green rays, which have a very strong tendency to
break up into spots, which are very rarely absent. The posterior slope is always
light in color, pale brown, reddish or yellowish, with hardly any green rays, so
that it is in sharp contrast to the rest of the shell.

The nacre is whitish, but has quite frequently salmon or pinkish tints, in fact,
delicate reddish tints are the rule in this variety, while the bluish-white of the
western marginata is rarely present.

Finally the posterior truncation, although similar to that of many specimens
of the western form, does not exhibit the extreme development, which is so often
seen in the latter.

Marshall, for instance, cites the Chemung and Tioga Rivers, belonging to the Susquehanna-
drainage. The Carnegie Museum possesses specimens from the Tioughnioga River, Susquehanna-
drainage, which surely are not typical marginata, but belong to the two eastern forms, var. susquehannce,
and the species varicosa.

188

MEMOIRS OF THE CARNEGIE MUSEUM.

Otherwise this form is like the western marginata, and differs markedly from
the common eastern A. varicosa, with which it is sometimes found associated.

Distance of beaks

Pr.ct. of

L.

H.

D.

from anterior end.

length.

1. Carlisle, Cat. No. 61.4677 (9 gravid). . . .

79 mm.

41 mm.

30 mm.

23 mm.

.29

2. do. “ “ do. (ci’)

.71 “

39 “

28 “

22 “

.30

3. do. “ “ do. ( 9 gravid) . . .

.69 “

36 “

29 “

23 “

.33

4. Selinsgrove, Cat. No. 61.4679 (9 gravid)

.66 “

35 “

26 “

19 “

.29

5. do. “ " do. (cT)

.61 “

36 “

28 “

20 “

.33

6. do. “ “ do. (cf)

.47 “

26 “

17 “

14.5“

.31

Soft parts and glochidia identical with those of A. marginata.

Breeding seaso7i: The following records for gravid females are at hand: Aug.
13, 1908; Aug. 13, 1910; Aug. 14, 1910; Aug. 20, 1909; Aug. 20, 1909; Aug. 21,
1909; Sept. 8, 1909.

These dates indicate only the beginning of the breeding season early in August
(eggs and glochidia found on first date!). Thus it is probably bradytictic.

Remarks: This is a new form, which has not been’ distinguished previously.
It comes very near to the western A. marginata in shape, having a rather sharp
])osterior ridge and a more or less distinctly truncated iiosterior slope, and having
the beaks nearer to the middle of the shell (at about one third of the length of
the shell). It is practically a marginata in shape with a peculiar color, and is thus
more closely allied to the western shell than to the eastern A. varicosa, and its
presence on the Atlantic side is highly interesting.

Wherever this form is found associated with the eastern varicosa, the two may
be distinguished at a glance, varicosa being smaller, with posterior ridge and pos-
terior truncation less developed, and with generally a darker epidermis, without
distinct spots, and the posterior slope not lighter, but rather darker than the rest
of the shell.

Localities represented in the Carnegie Museum:

Susquehanna River, Duncannon, Perry Co., Pennsylvania.

Susquehanna River, Selinsgrove, Snyder Co., Pennsylvania.

Conodoguinet Creek, Carlisle, Cumberland Co., Pennsylvania.

Juniata River, Juniata Bridge, Perry Co., Pennsylvania.

Frankstown Branch Juniata River, Huntingdon and Alexandria, Huntingdon Co., Pennsylvania.

(D. A. Atldnson).

Raystown Branch Juniata River, Ardenheim, Huntingdon Co.; and Mount Dallas, Bedford C5., Penn,
sylvania.

Penns Creek, Selinsgrove, Snyder Co., Pennsylvania.

North Branch Susquehanna River, Tunkhannock, Wyoming Co., Pennsylvania.

ORTMANN: monograph of the naiades of PENNSYLVANIA,

189

Chemung River, South Waverly, Bradford Co., Pennsylvania.

Tioughnioga River, Cortland, Cortland Co., New York (H. H. Smith) (drainage of upper North Branch
Susquehanna).

Localities represented in the Philadelphia Academy of Natural Sciences:

Juniata River, Tuscarora, Juniata Co. (S. N. Rhoads); Newton-Hamilton, Mifflin Co. (H. T. Mather,
Jr.); Mount Union, Huntingdon Co., Pennsylvania (S. N. Rhoads).

Distribution and Ecology (See figs. 18 & 19) : Type locality, Susquehanna
River, Selinsgrove, Snyder Co., Pennsylvania. Type set: Carnegie Museum Cat.
No. 61.4679.

The above localities are all that are known, and they are restricted to the
Susquehanna-drainage in Pennsylvania and New York. The form goes up the
North Branch and its tributaries, reaching at least to Cortland, New York, and
down the Susquehanna as far as the vicinity of Harrisburg (Duncannon). In
addition, it is found in Conodoguinet Creek and the Juniata River. In the latter
it ascends rather far (Mt. Dallas, Bedford Co.).

It should be noted, that it has not yet been found in the West Branch of the
Susquehanna; the few individuals at hand from the West Branch-drainage are

+ Alasmidonta marginata susquehannce.
m Alasmidonta varicosa.

all A, varicosa. However, this section of the Susquehanna system is poorly known,
and most of it is badly polluted by mine-water and refuse from tanneries. Below
Harrisburg, in the part of the Susquehanna which traverses the Piedmont Plateau,
and in its tributaries in that region, this form has not been observed.

The ecological habits of this variety are identical with those of the western

190

MEMOIRS OP THE CARNEGIE MUSEUM.

marginata. It frequents riffles, and this renders it the more conspicuous, since
there are few species on the Atlantic side which prefer this habitat.

According to our present , knowledge, A. marginata susquehannce is most
abundant in the Juniata River, and those parts of the Susquehanna system, which
are within the Allegheny Mountains. There is no trace of it in the Delaware or
Potomac drainages.

The peculiar features of the distribution suggest the idea, that this form came
from the West by crossing the Alleghenian divide. This has been discussed else-
where (Ortmann, 1913a, pp. 370 et seq).

Alasmidonta (Decurambis) varicosa (Lamarck) (1819).
Alasmidonta marginata Simpson, 1900, p. 670; Alas7nidonta varicosa (Lamarck)
PiLSBRY, 1901, p. 17; Alasmidonta varicosa (Lamarck) Simpson, 1914, p. 506.

Plate XII, fig. 5.

Records from Pennsylvania:

Lamarck, 1819 (Schuylkill River, Philadelpliia).

Lea, Obs. II, 1838, p. 56 (Crum Creek, Delaware Co.)^^^

Bruckhart, 1869 (Lancaster Co.).

Hartman & Michener, 1874 (Chester Co.)

Marshall, 1895 (Philadelphia).

Schick, 1895 (Tohickon Creek, Bucks Co.; Neshaminy Creek, Bucks Co.; Munckinipattus Creek,
Glenolden, Delaware Co.)

Ortmann, 19095, p. 207.

Caffrey, 1911 (Delaware River, Northampton Co.)

Characters of the shell: The shell of this species differs from that of A. marginata
by its smaller size, and by the outline, which is rather more subovate than sub-
trapezoidal, and slightly more elongate on the average. This difference of the
outline is brought about by a different development of the posterior ridge and
posterior slope. The posterior ridge is not so sharp, more broadly rounded, and
often appears biangulate,^^^ and the posterior slope is not so distinctly truncate.
This renders the whole posterior section of the shell more elongated and rounded,
and in consequence the anterior section is comparatively less developed, so that
the beaks are situated more anteriorly, generally at a distance from the anterior
end of considerably less than one-third of the length of the shell.

The color of the epidermis is much like that of A. marginata, with the darker

By a mistake I have given this record (Margaritana marginata) under Strophitus undulatus
(Ortmann, 19095, p. 205).

Indications of the biangulate character is sometimes found also in the western A. marginata.

OKTMANN: MONOGEAPH of the naiades op PENNSYLVANIA. 191

green shades prevailing. There is sometimes a light (brownish) ground-color,
with the rays not very strongly marked, but generally the dark green rays prevail,
and often the whole shell becomes dark green or blackish. The rays do not possess
a strong tendency to break up into spots, and are generally straight and simple,
but of various widths. Distinct spots are rarely seen. The posterior slope is
rarely lighter than the rest of the shell, and if so, the contrast is not well-marked;
but, on the contrary, the posterior slope often appears slightly darker than the
rest of the shell, which is due to a stronger development of dark green rays upon it.
Nacre whitish or bluish-white, very often with salmon, pinkish, or purplish shades,
which, however, are not very intense. Otherwise the shell is like that of A.
marginata.

The female shells are slightly more swollen in the region of the posterior ridge.
In old shells, this swelling is sometimes very distinct, and goes so far, that the
lateral faces of the shell, in front of the ridge, appear flat or even concave, which
concavity corresponds to a slight emargination of the lower margin. However,
sometimes males also show a slightly concave margin, and in many cases the sex
cannot be positively recognized by the characters of the shell, since not all females
have that swelling, and since the latter is sometimes also present in males.

Distance of beaks Pr.ct. of

L.

H.

D.

from anterior end.

length.

1.

Ridley Creek, Cat. No. 61.4680 (cf) ....

.70 mm.

38 mm.

29

mm.

18 mm.

.26

2.

Kunkletown, Cat. No. 61.4671 (9)

.69 “

36

((

26

a

17 “

.25

3.

do. “ “ do. (&)

.62 “

36

((

22

a

17 “

.27

4.

Greencastle, Cat. No. 61.4675 (c?)'

.61 “

33

i(

24

a

17.5“

.29

5.

Carlisle, Cat. No. 61.5873 (9 gravid). . .

.55 "

31

iC

22

a

14 “

.25

6. Mt. Dallas, Cat. No. 61.4266 (9 gravid)

.45 “

26

H

18

i(

12 “

.27

7.

Mantz, Cat. No. 61.4253 (cT)

.45 “

25

ii

18

((

12 “

.27

8.

do. “ “ do. ( 9 gravid) ....

.36 “

23

(C

15

<(

10 “

.28

There is another specimen from Ridley Creek, which is even larger than No.
1 (L. 75 mm.), but it is greatly deformed and freakish in shape, so that the dimen-
sions are entirely abnormal.

The above figures show the more anterior location of the beaks, although
the highest value corresponds to the lowest in A. marginata.

Soft parts and Glochidia identical with those of A. marginata.

Breeding season: Gravid females have been found: Aug. 9, 1910; Aug. 10,
1910; Aug. 11, 1910; Aug. 12, 1910; Aug. 13, 1908; Aug. 13, 1910; Aug. 14,
1910; Sept. 4, 1909; Sept. 5, 1909; Sept. 6, 1909; Sept. 8, 1909; and May 3,
1909.

The concave lower margin is here decidedly more frequent than in A. marginata.

192

MEMOIRS OF THE CARNEGIE MUSEUM.

Breeding begins in August, when eggs are present; later, in September, glo-
chidia are found. These are carried over the winter, and are still in the marsupium
in May; discharge has been observed on May 3.

This species supplements somewhat the incomiilete observations on A. mar-
ginata, and probably the breeding seasons of both species are about identical.

Remarks: This species has been frequently misunderstood, and confused with
the western A. marginata. But Simpson (1900) (following Wright) recognized
its specific distinctness, and I now hold the same opinion, although formerly
(Ortmann, 19096) I regarded it as a variety of A. marginata. But this was before
I had realized that there is in the Susquehanna-drainage another form, which
actually is more closely related to A. marginata.

A. varicosa is easily distinguished from A. marginata by the size and shape
of the shell, and from the var. marginata susquehannce it also differs in color. It is
generally much smaller than either of them, and specimens over 60 or 65 mm. in
length are quite scarce.

Although there is much variation in shape and color, this species hardly ever
inclines toward the western form, and no intergrades are found. I possess only a
single individual (dead shell) from White Clay Creek, Avondale, Chester Co.,
which has a sharper posterior ridge, and a lighter posterior slope, thus inclining
toward the var. susquehannce, but it has the general outline of varicosa, and, like
this, no spots. Being the only individual found at that locality, we cannot draw
any conclusions from it.

In the Susquehanna-drainage, I repeatedly found this species associated with
A. marginata susquehannce, but had never any difficulty in separating them. That
there are two distinguishable forms in this region, was evident to me when I col-
lected them together for the first time (Ardenheim, Aug. 13, 1908), and further -
study has only confirmed this view, although for a time I followed the views of
previous authors.

Localities in Pennsylvania, represented in the Carnegie Museum:

Delawar e-drainage :

Delaware River, Shawnee, Monroe Co.; Northampton Co. (G. W. Caffrey coll., G. H. Clapp donor);

Yardley, Bucks Co.

White Clay Creek, Avondale, Chester Co.

Ridley Creek, Delaware Co. (C. H. Conner).

Lehigh River, Bethlehem, Northampton Co. (Holland collection).

Princess Creek, Kunkletown, Monroe Co.

Lizard Creek, Mantz, Schuylkill Co.

Mahoning Creek, Lehighton, Carbon Co.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

193

Susquehanna-drainage:

Susquehanna River, York Haven, York Co.; Selinsgrove, Snyder Co.

Conewago Creek, Table Rock, Adams Co.

Conodoguinet Creek, Carlisle, Cumberland Co

Raystown Branch Juniata River, Ardenheim, Huntingdon Co.; Everett and Mount Dallas, Bedford Co.;

Bedford, Bedford Co. (A. Koenig).

Driftwood Branch, Sinnemahoning Creek, Driftwood, Cameron Co.

Cush Cushion Creek, Green Township, Indiana Co. (D. A. Atkinson).

Potomac-drainage :

Conococheague Creek, Greencastle and Scotland, Franklin Co.

West Branch Conococheague Creek, Mercersburg Junction, Franklin Co.

Great Tonoloway Creek, Thompson Township, Fulton Co.

Localities represented in the Philadelphia Academy of Natural Sciences:

Delaware River, Delaware Water Gap, Monroe Co. (S. N. Rhoads), and Columbia, Warren Co., New
Jersey (S. N. Rhoads).

Big Neshaminy Creek, Edderton,i^^ Bucks Co. (H. W. Fowler).

Pennypack Creek, Holmesburg, Philadelphia Co. (H. W. Fowler).

Swamp Creek, Zieglerville, Montgomery Co. (Bayard Long).

Manatawny Creek, Earlville, Berks Co. (H. A. Pilsbry).

Maiden Creek, Berks Co.

Sacony Creek, Kutztown, Berks Co. (H. K. Deisher).

Lancaster, Lancaster Co. (J. B. Hshleman).

Susquehanna River, York Furnace, York Co. (W. Stone).

Other localities represented in the Carnegie Museum:

Stony Brook, Princeton, Mercer Co., New Jersey.

Tioughnioga River, Cortland, Cortland Co., New York (H. H. Smith).

Potomac River, Hancock, Washington Co., Maryland.

Wills Creek, Ellerslie, Allegany Co., Maryland.

South Branch Potomac River, Southbranch, Hampshire Co., West Virginia.

Shenandoah River, Harpers Ferry, Jefferson Co., West Virginia.

South Fork Shenandoah River, Elkton, Rockingham Co., Virginia.

South River, Waynesboro, Augusta Co., Virginia. (Headwaters of South Fork of Shenandoah.)
Catawba River, Bridgewater, Burke Co., North Carolina.

Distribution and Ecology in Pennsylvania (See fig. 19): From the published
records it is seen that this species belongs to the Atlantic-drainage in Pennsylvania,
and that it is rather evenly distributed over it, with the possible exception of
the larger rivers. Specimens found by myself in the Delaware and lower Susque-
hanna are only few. In the smaller streams this species is more abundant, and
locally common. This agrees well with the conditions seen in the western A.
marginata, and with this A. varicosa also agrees in that it is distinctly a shell which

Probably Eddington.

194

MEMOIRS OF THE CARNEGIE MUSEUM.

prefers strong currents and gravelly bottoms, thus being most frequently found
in and near riffles. In the mountains it goes far up into the headwaters, and the
most western locality is in Indiana Co., in a tributary of the West Branch of the
Susquehanna (Cush Cushion Creek).

General distribution. Type locality, Schuylkill River, Philadelphia (Lamarck).

According to Simpson (1900), this species extends over the Atlantic-drainage
from the lower St. Lawi*ence to South Carolina.

As has been stated above, the mutual relations of A. varicosa and marginata
in central and western New York are doubtful, and specimens from New York
are generally recorded under the name of marginata, although it is beyond doubt,
according to our material, that varicosa is found in the upper Susquehanna-drainage
in New York, and, according to the figures of DeKay in other parts of the state also.

The species has been reported from Maine (Lermond, 1909), New Hampshire
(Call), Massachusetts (Gould-Binney, 1870), Rhode Island (Carpenter, 1890),
and Connecticut (Linsley, 1845) but Johnson (1915, p. 27) erroneously calls the
New England form A. ^narginata. Southward from Pennsylvania, A. varicosa is
known from Red Clay Creek, Christiania Township, Newcastle Co., Delaware
(Rhoads, 1904), and from the Potomac-drainage, in addition to the localities repre-
sented in the Carnegie Museum, from Sideling Creek, Allegany Co., Maryland,
the Potomac River, Cherry Run, jMorgan Co., West Virginia (Pilsbry, 1894).

Farther South, exact localities are missing, but Simpson’s statement, that
it extends to South Carolina, is substantiated by specimens collected by myself in
the upper Catawba River in North Carolina. Among the latter are individuals
which fully agree with the Pennsylvanian form, but this set is extremely variable
in shape and especially in color.

A. varicosa of the Atlantic slope is the representant of the western A. mar-
ginata, and in its distribution apparently falls in line with several other Atlantic
species, constituting a northern stock in the fauna of the Atlantic slope. This
has been discussed elsewhere (Ortmann, 1913a, p. 363 & 370).

Walker (1913, p. 22) reports this species from Lake Erie; this, however, re-
quires renewed attention, since the form from Lake Erie may belong to A. mar-
ginata (see above). I have never seen the lake-form, and thus I am unable to
judge.

ORTMANN; monograph of the naiades of PENNSYLVANIA. 195

Genus Strophitus Rafinesque (1820).

Ortmann, 1912, p. 299; Simpson, 1914, p. 344.

Type Anodonta undulata Say.

Two species of this genus are generally credited to Pennsylvania, but I have
been able to find only one {S. edentulus), which is distributed all over the state
both on the western and the eastern side of the mountains. The type-species
possibly originally came from Pennsylvania, but being founded upon a very poor
and immature specimen, has been largely misunderstood. Simpson (1900, p. 618;
1914, p. 345 & 349) admits the existence, on the Atlantic side, of “a small, thin
form, . . . usually biangulate behind,” but I have not been able to locate it.

Conner believes, that he has rediscovered the true S. undulatus of Say, and I
have received from him specimens from the New Jersey side of the Delaware River
(not far from the supposed type-locality of Say’s specimen, on the Pennsylvania
side of the river) ^ However, I can see only a local race of the common S. edentulus
in this, probably belonging to the tidewater region of the Delaware River, and
connected by intergrades with the common form, wherever the latter goes into
large and more quiet bodies of water. But since my material is entirely insufficient
to solve the question, I shall mention here the two forms as species, thus avoiding
the inconvenience of having to call the typical, common form by a varietal name
(*S. undulatus edentidus) , and making the local race the main species {S. undulatus) .

Strophitus undulatus (Say) (1817).^®^

Simpson, 1914, p. 349 (but synonymy largely incorrect).

Plate XII, fig. 6.

Records from Pennsylvania :

Say does not give a type-locality for his Anodonta undulata, but his specimen probably came from
near Philadelphia.

Lea (Obs. II, 1838, p. 54, and Obs. X, 1863, p. 450) repeatedly reports A. undulata Say from the
Schuylkill River, near Philadelphia, and distinguishes it from A. edentula. The mouth of the Schuylkill,
on the Delaware is just opposite the locality where Conner obtained liis specimens.

All other records for this species are either very doubtful, whether they belong to the true undu-
latus, or are positively referable to S. edentulus.

(See Ortmann, 19096, pp. 194 & 205.)

Characters of the species: Like S. edentulus in every particular, except that the
beaks are slightly more inflated, elevated, and incurved. In consequence of this
the shell appears higher anteriorly, and more tapering and pointed posteriorly.

136 Not 1816.

196

MEMOIRS OF THE CARNEGIE MUSEUM.

L. H. D.

Size: 1. ( 9 ) 53 mm. 33 mm. 23 mm.

2 52 “ 33 “ 24 “

These are the two largest at hand.

Soft parts: I have seen only those of a sterile female, and find them identical
with those of S. edentulus. Glochidia unknown.

Breeding season: Conner (1909, p. 112, as Anodonta undulata) found this
form gravid in December and IMarch, but not in April and May.

Remarks: For all the material I possess I am indebted to the kindness of Mr.
C. H. Conner, who sent me eight specimens, all from the tide-waters of the Dela-
ware River, near Newbold, Gloucester Co., New Jersey. These specimens show
the differential characters given above. ]\Ir. L. S. Frierson {in litteris) has called
my attention to certain supposed differences in the hinge, but I am unable to see
them, hloreover the shape of the shell is somewhat variable in the eight specimens
at hand, and further I have specimens of the western S, edentulus (chiefly from quiet
waters) which come very near to this form from the Delaware in the inflation of the
beaks. Further I have certain large and old specimens of S. edentulus, where the
inflation of the beaks is much greater than in any specimen from the Delaware.
For instance a giant 'Specimen from Cush Cushion Creek (abandoned reservoir)
is very remarkable in this respect; but it was found among a large number of
typical edentulus and cannot be anything else.

In the hinge there is the same variation as in S. edentulus. In some speci-
mens vestiges of teeth are seen, in others they are almost entirely obliterated, and
indicated only by slight thickenings and curves of the hinge-line. Mr. Frierson
maintains that the tooth of the left valve (cardinal -|- interdental) is absent, and
reiiresented by an excavation. This does not hold good for all specimens before
me, and, on the other hand, this is a condition, very often seen in S. edentulus.

I am very much inclined to think that this is only a local variety, or even
only an ecological race, belonging to the tidewaters of the Delaware River, and
localities with a similar environment. In the Delaware, just below Trenton,
New Jersey (at Penns Manor, Bucks Co., Pennsylvania) I collected a specimen,
which is rather small, and well agrees in outline with the specimens received from
Mr. Conner, but the beaks are not inflated and elevated. Specimens from the
Delaware River, Trenton, in the Philadelphia Academy, are typical S. edentulus.
On the other hand, a specimen of S. edentulus from the Delaware-Raritan Canal,
specimens from Conneaut Lake, Lake Erie, and Winona Lake, have the shells
inflated more than usual, although the beaks are not so elevated as in the form
from the lower Delaware, and thus the outline is more regularly elliptical, and

ORTMANN: monograph of the naiades of PENNSYLVANIA.

197

not ovate (higher anteriorly). Then again the ovate outline, with the anterior
part of the shell high, and the posterior part more or less tapering, is also repre-
sented in typical S. edentulus with compressed shell.

The real S. undulatus therefore remains a disputed form.

Of course S. undulatus, as discussed here, is not fully identical with Simpson’s
undulatus, for Simpson included in this all small, thin, and generally distinctly
biangulate shells (Simpson, 1900, p. 618, footnote). But these characters well fit
young specimens of S. edentulus, while the specimens received from Conner are not
biangulate behind, on account of the tapering posterior end.

Locality represented in the Carnegie Museum:

Delaware River, Newbold, Gloucester Co., New Jersey (C. H. Conner).

Distribution (See fig. 21) : Frierson has sent me for examination a specimen
from Warwick Pond, Warwick, Rhode Island, and another one from Orange Co.,
Virginia, but I have not been able to satisfy myself that they belong here, as was
supposed, and that they are a species distinct from S. edentulus. Johnson (1915)
keeps S. undulatus apart from S. edentulus, and gives for the former, a number of
localities in New England, but I have no means of deciding what he understood
by this species. No other records are known positively referring to this form.

Strophitus EDENTULUS (Say) (1829).

Strophitus edentulus (Say) Simpson, 1914, p. 345, and Strophitus undulatus Simpson
{pro parte), ibid., p. 349.

Plate XII, figs. 7, 8.

Records from Pennsylvania:

? Lea, Obs. X, 1863, p. 450 (Schuylkill River, Philadelphia).'^®

Bruckhart, 1869 (as edentula) (Lancaster Co.)

Hartman & Michener, 1874 (as edentula) (Brandywine Creek, Chester Co.)

Harn, 1891 (as undulata) (western Pennsylvania).

Scliick, 1895 (as undulata) (Canal, 27th ward, Philadelphia).

Marshall, 1895 (as edentula) (Allegheny River, Warren Co.)

Rhoads, 1899 (as edentula) (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.; Beaver
River, Wampum, Lawrence Co.)

Ortmann, 19096 (as undulatus), p. 194, 202, 207.

Characters of the shell: Shell of medium size, rather thin when young, moder-
ately thick when old. Outline subelliptical to subovate, broadly rounded in front,
more narrowly rounded behind, sometimes somewhat pointed or biangulate behind.

Lea frequently refers to A. undulata as coming from the Schuylkill River. He distinguishes it
from A. edentula, but we cannot tell which form he had in mind.

198

MEMOIRS OF THE CARNEGIE MUSEUM.

Lower margin convex or straight, sometimes slightly concave in the middle. Upper
margin gently convex, forming a more or less distinct angle with the obliquely
descending posterior margin. Beaks little prominent, situated in front of the
middle. Beak-sculpture distinct, moderately heavy, consisting of about four
concentric bars, placed obliquely with reference to the hinge line, rounded anter-
iorly, angular upon the posterior ridge. A few radiating lines anteriorly and
posteriorly to the bars (See Marshall, 1890, fig. 12).

Shell generally gently convex or rather flat upon the sides, but occasionally
more or less swollen or inflated. Posterior ridge indistinct and broadly rounded.
Posterior slope somewhat compressed, and, when young, often elevated at the
upper posterior angle.

Epidermis yellowish, greenish, brownish, or blackish, with more or less distinct
greenish or blackish rays. Rays straight, narrow or wider, often quite obsolete.
Posterior slope sometimes darker than the rest of the shell. Very old shells are
mostly quite black all over. Concentric darker bands may be present.

Hinge much reduced. Lateral teeth entirely absent; pseudocardinals indi-
cated only by slight swellings of the hinge-line; rarely can the single elements be
made out. Generally, the hinge-line is only somewhat wavy under the beaks.
Beak-cavity moderate. Dorsal muscle-scars in the beak-cavity. Anterior ad-
ductor-scars rather distinct, posterior ones less so. Nacre bluish white, toward the
thick part of the shell pure white, or cream-color, and very often salmon-pinkish.

]\'Iale and female shells practically indistinguishable.

L. H. D.

Size: 1. Cush Cushion Creek, Cat. No. 61.3627 (d^) 106 mm. 56 mm. 43 mm.

2. Grove City, Cat. No. 61.3278 ($ gravid) 102 “ 55 “ 39 “

3. Tioga, Cat. No. 61.4154 (9 gravid) 81 “ 41 “ 32 “

4. Milesburg, Cat. No. 61.4158 (9 gravid) 74 “ 42 “ 27 “

5. Warren, Cat. No. 61.4164 (d") 66 “ 37 “ 22 “

6. Darlington, Cat. No. 61.2945 (cf ) 44 “ 25 “ 16 "

Soft parts: A poor figure was published by Lea (Obs. II, 1838, PI. 15, fig. 47) ;
a description was given by Ortmann, 1912, p. 299. Glochidia figured by Lea
(Obs. VI, 1858, PL 5, fig. 5), and Surber (1912, PL 1, fig. 3). Siirber’s measure-
ments are: 0.350 X 0.285, while mine {1. c., p. 300) are: 0.36 X 0.30 mm.

Breeding season: Records for gravid females are most complete. They cover
the time from July 11 to November 4, and from April 17 to May 22. It is a typi-
cally hradytictic form, with the interim between the breeding seasons falling,
in our region, in the end of May, June, and the beginning of July. Surber (1912)
gives a shorter interval, only in July.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

199

Remarks: Although very variable in outline and color, this species is easily
recognized by the elliptical shape, and the rudimentary condition of the hinge.
If gravid females are at hand, the marsupial structure unmistakably indicates the
genus. Only Anodontoides ferussacianus might be confounded with this species,
but the shell of the latter is generally thinner, and the nacre is more bluish white,
with very little pure white, and further the beak-sculpture is much finer.

Specimens with well-defined rays have been called pavonia Lea. The type
locality of this is Little Beaver Creek in Ohio. I possess many specimens from
this creek in Pennsylvania, and among them are many which correspond to pavonia,
but they are connected by all possible intergrades with the normal S. edentulus.

• Strophitus edentulus (western half of State of Pennsylvania).

Such brightly colored individuals may turn up anywhere, on the western as well
as on the eastern side of the Alleghenies, and are always found associated with
the normal form.’^®’^

There is little inclination to form local races, although under certain conditions
and at certain localities the specimens look rather uniform. Thus in the mountain-
streams on the western side of the mountains, there is a small, often more elongated
race of S. edentulus, and the Allegheny River in general, does not produce very
large, but rather heavy-shelled specimens. In the Beaver-drainage this species
is much larger on the average. Strangely enough the largest specimens on record
are from the Atlantic-drainage, in the headwaters of the West Branch of the

Wilson & Clark (1912a, p. 48) believe that the better development of rays (in pavonia) is corre-
lated with the clearness of the water, and I think that this is right.

200

MEMOIRS OF THE CARNEGIE MUSEUM.

Susquehanna (Cush Cushion Creek) ; but these came from an old, abandoned
reservoir, with a muddy bottom and rich vegetation.

Specimens from Conneaut Creek, tributary to Lake Erie, do not differ from
the normal type. However, in Lake Erie proper a peculiar form is found. I my-
self did not find many specimens, but I have received additional ones from C.
Goodrich, and all these represent a dwarfed race (largest at hand: L. 51 mm..

• Strophiius edentvlus (eastern half).
+ Strophitus undulatus.

H. 28 mm., D. 20 mm.), of a rather regular, subelliptical and elongate outline.
I do not, however, think it advisable to distinguish this race by a varietal name.

Localities in Pennsylvania, represented in the Carnegie Museum:

Small tributaries of the Ohio:

Buffalo Creek, Acheson, Washington Co.

Cross Creek, Avella, Washington Co.

Little Beaver Creek, Cannelton (H. H. Smith), Darlington, New Galilee, Beaver Co.; Enon Valley,
Lawrence Co.

Raccoon Creek, Raccoon Township and New Sheffield, Beaver Co.; Bavington, Washington Co.
Chartiers Creek, Carnegie, Allegheny Co. (D. A. Atkinson & J. L. Graf).

Little Chartiers Creek, Morganza, Washington Co.

Beaver-drainage :

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Ellwood City, Lawrence Co. (G. H. Clapp & H. H. Smith); Celia, Beaver Co.;

Zelienople, Butler Co.; Harmony, Butler Co. (R. G. Pflaum).

Slipperyrock Creek, Wurtemberg and Rose Point, Lawrence Co.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

201

Wolf Creek, Grove City, Mercer Co.

Brush Creek, Celia, Beaver Co.

Little Connoquenessing Creek, Harmony, Butler Co.

Thorn Creek, McBride, Butler Co.

Bonnie Brook, East Butler, Butler Co.

Mahoning River, Mahoningtown, Coverts, and Edinburg, Lawrence Co.

Neshannock Creek, Eastbrook, and Volant, Lawrence Co.; Leesburg, Mercer Co.

Otter Creek, Mercer, Mercer Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Shenango and Jamestown, Mercer Co.;
Linnesville, Crawford Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

A llegheny-drainage :

Allegheny River, Natrona, Allegheny Co.; Aladdin, Godfrey, Kelly, Mosgrove, and Templeton, Armstrong
Co.; Hickory, Forest Co.; Warren, Warren Co.

Buffalo Creek, Harbison, Butler Co.

Loyalhanha River, Idlepark and Ligonier, Westmoreland Co.

Beaver Run, Delmont, Westmoreland Co.

Blacklegs Creek, Saltsburg, Indiana Co.

Yellow Creek, Homer, Indiana Co.

Quemahoning Creek, Stanton’s Mill, Somerset Co.

Crooked Creek, Rosston, Armstrong Co.; Creekside, Indiana Co.

Little Mahoning Creek, Goodville, Indiana Co.

Sandy Creek, Sandylake, Mercer Co.

French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co.
Conneaut Outlet, Conneautlake, Crawford Co.

Conneaut Lake, Crawford Co.

Conheauttee Creek, Edinboro, Erie Co.

Brokenstraw Creek, Garland, Warren Co.

Connewango Creek, Russell, Warren Co.

Potato Creek, Smethport, McKean Co. (P. E. Nordgren).

Monongahela-drainage :

Monongahela River, Elizabeth, Allegheny Co. (D. A. Atkinson).

Youghiogheny River, Confluence, Somerset Co.

Tenmile Creek, Clarksville, Greene Co.; Amity, Washington Co.

South Fork Tenmile Creek, Waynesburg, Greene Co.

Dunkard Creek, Wiley and Mount Morris, Greene Co.

Cheat River, Cheat- Haven, Fayette Co.

Lake Erie-drainage :

Conneaut Creek, West Springfield, Erie Co.; Springboro, Crawford Co.

Lake Erie, Presque Isle Bay, Erie, Erie Co.

Genesee River-drainage:

Genesee River, Genesee, Potter Co.

202

MEMOIRS OF THE CARNEGIE MUSEUM.

Delaware-drainage: ■> - .

Delaware River, Penns Manor and Yardley, Bucks Co.; Shawnee, Monroe Co.

Schuylkill River, Manayunk, Philadelphia Co.

Schuylkill Canal, Manayunk, Philadelphia Co.

Wissahickon Creek, Roxboro, Philadelphia Co.

Little Neshaminy Creek, Grenoble, Bucks Co.

Princess Creek, Kunkletown, Monroe Co.

Lizard Creek, Mantz, Schuylkill Co.

Susquehanna-drainage :

Susquehanna River, Selinsgrove, Snyder Co.

Conewago Creek, York Haven, York Co.; Table Rock, Adams Co.

Conodoguinet Creek, Carlisle, Cumberland Co.

Cocolamus Creek, Cocolamus, Juniata Co. (D. A. Atkinson).

Lost Creek, Mifflintown, Juniata Co. (D. A. Atkinson).

Raystown Branch Juniata River, Everett and Mount Dallas, Bedford Co.

Dunning Creek, Bedford, Bedford Co. (J. F. L. Raschen).

West Branch Mahantango Creek, Richfield, Juniata Co.

Middle Creek, Freeburg, Snyder Co. (D. A. Atkinson).

Bald Eagle Creek, Milesburg, Center Co.

Beaver Dam Creek, Flinton, Cambria Co. (D. A. Atkinson).

Swartz Run, Ashville, Cambria Co.

Chest Creek, Patton, Cambria Co.

Cush Cushion Creek, Green Township, Indiana Co.

North Branch Susquehanna River, Tunkhannock, Wyoming Co.

Chemung River, South Waverljq Bradford Co.

Millrace of Crooked Creek, Tioga, Tioga Co.

Potomac-drainage :

East Branch Little Antietam Creek, Waynesboro, Franklin Co.

Conococheague Creek, Greencastle and Scotland, Franklin Co.

West Branch Conococheague Creek, Mercersburg Junction, Franklin Co.

Pennsylvanian localities represented in the Philadelphia Academy of Natural
Sciences:

Big Elk Creek, Westgrove, Chester Co. (C. H. Conner)

Chester Creek, Delaware Co.

Maiden Creek, Berks Co.

Big Neshaminy Creek, “ Edderton ” (probably Eddington), Bucks Co. (H. W. Fowler).

Columbia, Lancaster Co. (J. B. Eshleman).

Little Swatara Creek, Jonestown, Lebanon Co. (C. H. Conner).

Additional locality in New Jersey:

Delaware River, Trenton, Mercer Co., New Jersey (H. Hamilton).

Not exact, either creek or town given incorrectly.

OKTMANN: monograph of the naiades of PENNSYLVANIA.

203

Other localities represented in the Carnegie Museum: ■

Lake-drainage:

Seneca Co., New York (Smith collection).

Conestogo River, Conestogo, Waterloo Co., Ontario, Canada (Miss Maria Jentsch).

Lake Erie, Port Dover, Norfolk Co., Ontario, Canada; Vermilion, Erie Co., Ohio; La Plaisance Bay
Monroe Co., Michigan (C. Goodrich).

Creek at North Fairfield, Huron Co., Ohio (0. E. Jennings).

Sandusky River, Upper Sandusky, Wyandot Co., Ohio (C. Goodrich).

Swan Creek and Ten Mile Creek, Toledo, Lucas Co., Ohio (C. Goodrich).

St. Marys River, Rockford, Mercer Co., Ohio (C. Goodrich).

Beaver Creek, Williams Co., Ohio (C. Goodrich).

Maumee River, Fort Wayne, Allen Co., Indiana (C. Goodrich).

Otter Creek, Monroe Co., Michigan (C. Goodrich).

Raisin River, Adrian and Tecumseh, Lenawee Co., Michigan (C. Goodrich).

Drainage of Red River of the North:

Sheyenne River, Argusville, Cass Co., North Dakota (S. M. Edwards).

Ohio-drainage: .

West Branch Nimishillen Creek, Canton, Stark Co., Ohio.

Tuscarawas River, Ohio (Holland collection).

Scioto River, Columbus, Franklin Co., Ohio (Smith collection).

Wabash River, St. Henry, Mercer Co., Ohio (C. Goodrich).

Winona Lake, Kosciusko Co., Indiana (E. B. Williamson).

Cheat River, Jaco and Mount Chateau, Monongalia Co., West Virginia.

Tygart River, Elkins, Randolph Co., West Virginia.

French Creek, Hampton, Upshur Co., West Virginia.

West Fork River, Lynch Mines, Harrison Co.; Weston, Lewis Co., West Virginia.

Little Kanawha River, Burnsville, Braxton Co., West Virginia.

North Fork Hughes River, Cornwallis, Llitchie Co., West Virginia.

Elk River, Shelton and Clay, Clay Co.; Gassaway and Sutton, Braxton Co., West Virginia.

Coal River, Sproul, Kanawha Co., West Virginia.

Licking River, Farmer, Rowan Co., Kentucky.

Cumberland- and Tennessee-drainages:

Cumberland River, Burnside, Pulaski Co., Kentucky (B. Walker, donor, soft parts only).

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).

Elk River, Estill Springs, Franklin Co., Tennessee (H. H. Smith).

Tennessee River, Concord, Knox Co., Tennessee.

French Broad River, Boyd Creek, Sevier Co., Tennessee.

Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill, Noeton, and
Holston Station, Grainger Co., Tennessee.

South Fork Holston River, Pactolus and Bluff City, Sullivan Co., Tennessee.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia; Saltville,
Smyth Co., Virginia.

Clinch River, Solway, Knox Co.; Edgemoor and Clinton, Anderson Co.; Clinch River Station, Claiborne
Co.; Oakman, Grainger Co., Tennessee; St. Paul, Wise Co., Cleveland, Russell Co., Raven, Rich-
land, and Cedar Bluff, Tazewell Co., Virginia.

204

MEMOIRS OF THE CARNEGIE MUSEUM.

Powell River, Combs, Claiborne Co., Tennessee; Dryden, Lee Co., Virginia.

South Fork Powell River, Big Stone Gap, Wise Co., Virginia.

Mississippi River and westward:

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection).

Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).

Hinkston Creek, Columbia, Boone Co., Missouri (D. K. Greger).

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

White River, Hollister, Taney Co., Missouri (W. I. Utterback) ; Cotter and Norfolk, Baxter Co., Arkansas
(A. A. Hinkley).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).

Saline River, Benton, Saline Co., Arkansas (H. E. Wheeler).

Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).

Lawrence, Douglas Co., Kansas (R. L. Moo die).

Fourteen Mile Creek, Fort Gibson, Muskogee Co., Oklahoma (F. B. Isely).

Atlantic-drainage :

Delaware-Raritan Canal, Princeton, Mercer Co., New Jersey.

Potomac River, Hancock, Washington Co., Maryland.

South Branch Potomac River, Romney, Hampshire Co., West Virginia.

North Fork Shenandoah River, Broadway, Rockingham Co., Virginia.

South Fork Shenandoah River, Elkton, Rockingham Co., Virginia.

South River, Wajmesboro, Augusta Co., Virginia.

Rappahannock River, Remington, Fauquier Co., Virginia.

Rapidan River, Rapidan, Culpeper Co., Virginia.

Mountain Run, Culpeper, Culpeper Co., Virginia.

North River, Buena Vista and Lexington, Rockbridge Co., Virginia.

Calf Pasture River, Goshen, Rockbridge Co., Virginia.

Roanoke River, Salem, Roanoke Co., Virginia.

Mason Creek, Salem, Roanoke Co., Virginia.

Distribution and Ecology in Pennsylvania (See figs. 20 & 21) : This is the
only species of Naiad whose range covers the whole of the state of Pennsylvania,
the divide of the Allegheny Mountains having no influence whatever upon it.
The forms found on either side of the divide do not differ at all, and go far up into
the headwaters, so that there is practical continuity of distribution across the
divide. There is no doubt that this distribution is to be accounted for by an
actual crossing over from the western to the eastern slope, and probably this has
been effected by stream piracy. This has been discussed elsewhere (Ortmann,
1913a, pp. 367-369).

Strophitus edentulus also is extremely common all over the state, although it
is distinctly averse to large rivers. Rhoads reports it from the Ohio below Pitts-
burgh, but I have never found it there. Above Pittsburgh it appears in the Alle-
gheny, but is decidedly scarce in Allegheny and Armstrong Cos., and the specimens

ORTMANN: monograph of the naiades of PENNSYLVANIA.

205

are rather smalLon the average. Farther up, and in the tributaries, it becomes
more and more abundant, and in all small creeks it is a common shell.

The fact is remarkable, that S. edentulus is the only Naiad present in the
upper Youghiogheny. There is no other stream in Pennsylvania, which offers a
parallel case, but such are found in several mountain streams in West Virginia.
This has been discussed elsewhere (Ortmann, 1913a, pp. 303, 305, 308; 357, 367
et seq.).

Likewise on the Atlantic slope this species is scarce or missing in the large
rivers. This is most evident in the lower Susquehanna. It is present in the
Delaware near and above Trenton, New Jersey, but does not seem to be abundant.
It may be that in the estuary of the Delaware S. undulatus takes its place.

We may therefore call 8. edentulus a form characteristic of smaller streams.
Baker (1898a, p. 68) says indeed that it is found in the larger lakes and rivers on
muddy bottoms, while Scammon (1906, p. 323) reports it to prefer mud and quiet
water, and to be most abundant in smaller streams (in Kansas). Its absence
in the larger rivers in Pennsylvania may be connected with its aversion to rough
bottom and strong currents. Even in the small streams it avoids riffles, but de-
lights in quiet and protected nooks, pools, and eddies, where there is a moderate
and rather uniform current, and a deposit of fine gravel, sand, or mud. It is often
found in the Dianther a-patches.

On the other hand this species goes into canals and lakes. Altogether, it
lives under a great variety of environmental conditions, and thus it is not aston-
ishing that it is so widely distributed.

General distribution: Type locality, Wabash River (Say).

According to Simpson (1900) 8. edentulus ranges over the “entire Mississippi-
drainage; the St. Lawrence system and south in streams draining into the Atlantic
to North Carolina; north in the British possessions to Lake Winnipeg; south-
west to central Texas.” (An Alabaman locality has also been recorded.)

This is substantially correct, and this is possibly the widest range occupied
by any species of Naiad in North America. Its occurrence in Canada in the lower
St. Lawrence-drainage, as well as in the west, is well-established (Whiteaves,
1863; Bell, 1859; Call, 1885; Dawson, 1875; Hanham, 1899; Christy, 1855; see
also our records from Ontario and North Dakota). Its presence on the Atlantic
slope from Maine (Lermond, 1909) southward is well known, and substantiated
by our records as far South as the Roanoke system in Virginia. It is said to go
to North Carolina (Simpson), but here the southern boundary is obscurely known.

The center of radiation is apparently located in the Mississippi system (See

206

MEMOIRS OF THE CARNEGIE MUSEUM.

Ortmann, 1913a, p. 367 et seq.). It is found everywhere in the drainage of the
Ohio and in that of the upper Mississippi. There are sections, where records are
scarce or missing, as for instance in Kentucky, and its absence in the upper Kanawha
system in West Virginia and Virginia should be noted (See Ortmann, 1913, pp.
308 and 367). It seems to be rare in the Cumberland (Wilson & Clark, 1914),
but in the upper Tennessee system in Tennessee and Virginia it is common.

In a westerly and southwesterly direction it goes across the Mississippi, and
has been found as far west as eastern Nebraska (Tryon, 1868); in Kansas, as
far as Saline Co., in the Kansas-drainage, and Reno County, in the southern
drainage (Scammon, 1906). From Arkansas it has been reported by Call (1895)
and Meek & Clark (1912), and it is represented from the Ozark streams, the Saline
and Ouachita Rivers in the Carnegie Museum (previously reported from Ouachita
by Vanatta, 1910, and Wheeler, 1918). It is also present in Oklahoma (Carnegie
Museum). Frierson (1899) reports it from a tributary of Sabine River in De Soto
Parish, Louisiana, and according to Simpson it goes to central Texas (Vaughan,
1893, does not mention it from northwestern Ijouisiana, and Singley, 1893, does
not mention it from Texas).

Records from the southern Atlantic and Gulf states are lacking, though
Simpson cites Tyner, Tuscaloosa Co., Alabama; and a number of allied forms,
separated as species, are credited to Georgia, Alabama, and Mississippi. I can-
not judge of these; but I may mention, that specimens from the Alabama-drainage
in the Carnegie Museum, partly collected by H. H. Smith, partly by myself, seem
to differ, and to represent possibly two other species: S. connasaugaensis (Lea)
(= alabamensis Lea = gesneri Lea), and S. spillmani (Lea). Both' have the
pseudocardinal teeth better developed than S. edentulus.

Subfamily LAVIPSILIN^F) (v. Ihering, 1901) Ortmann (1910).^^®
Ortmann, 1910, p. 118; 1911, p. 337; 1912, p. 300.

Key to the Genera of the Lampsilin^e.

ai. Marsupium not kidney-shaped. Ovisacs subcylindrical or very slightly compressed. Placentae
generally very solid. Inner edge of mantle in front of branchial opening, not distinctly differ-
entiated. Shell rounded, ovate, subelliptical, sometimes with sculpture upon the disk. Male
and female shells practically alike.

bi. Marsupium occupying the whole of the edge of the outer giU, folded. Placentae club-shaped,
short. Shell subelliptical, smooth Ellipsaria.

62. Marsupium occupying only a part of the outer gill. Placentae subcylindrical, elongated or very
long. Shell more or less rounded, generally with tubercles.

V. Ihering first coined tliis subfamily-name, but places some of the genera belonging here in his
subfamily Quadrulince.

if

OETMANN: MONOGEAPH op the naiades of PENNSYLVANIA. 207

Cl. Placentae moderately long, slightly curved. Marsupium just behind the middle of the gill.

Outside of shell with a few large knobs Obliquaria.

C2. Placentae very long, spirally coiled up. Marsupium in, or slightly in front of, the middle of

the gill. Outside of shell with numerous small tubercles Cyprogenia.

a 2. Marsupium kidney-shaped. Ovisacs lanceolate, dilated and compressed. Placentae not very solid.

Inner edge of mantle in front of branchial opening, more or less differentiated. Shell rounded,
elliptical, or elongate, without Sculpture on the disk.

61. Inner edge of mantle, in front of branchial opening, slightly lamellate and crenulated, but without

papillae or flaps. Male and female shell differing only slightly in shape, or hardly at all.
Cl. Shell rounded, ovate, or subelliptical. Glochidia of normal size and shape, subovate.

di. Shell rounded or short-ovate. Epidermis brownish, rarely greenish, with indistinct

rays Obovaria.

d2. Shell ovate or subelliptical, elongate. Epidermis greenish or yellowish, with more or

less distinct rays , Actinonaias.

C2. Shell ovate, triangular, or subelliptical. Glochidia either of abnormal size or shape.
di. Shell subovate or subtriangular, with a strong posterior ridge.

Cl. Glochidia of normal shape (subovate), but of abnormally small size. Shell sub-
ovate or elongate Amygdalonaias.

6 2. Glochidia spatulate, with gaping margins, very large. Shell subtriangular.

Plagiola.

d2. Shell subovate or subelliptical, generally compressed and winged at the upper posterior
part, without distinct posterior ridge.

Cl. Glochidia of normal shape, but of abnormally small size. Shell rather thin.

Paraptera.

6 2. Glochidia celt-shaped, with two spines on each valve. Shell thicker Propter a.

62. Inner edge of mantle, in front of branchial opening, with papillae or flaps. Male and female

shell distinctly, and often greatly, different in shape.

Cl. Inner edge of mantle parallel with and close to the outer edge. Glochidia subovate or
subelliptical.

di. Inner edge of mantle with papillae.

Cl. Papillae few, represented chiefly by one large caruncle. Shell very small. Beak-

sculpture concentric Toxolasma.

62. Papillae more numerous. Shell small, medium, or large. Beak-sculpture double-

looped Eurynia.

d2. Inner edge of mantle forming a ribbon-like flap, projecting at the anterior end. Shell

large Lampsilis.

C2. Inner edge of mantle, in front of branchial opening, more or less remote from outer edge in
the female. Glochidia subcircular. Shell rather small, of peculiar and various shapes,
chiefly in the female Truncilla.

Genus Ellipsaeia Rafinesque (1820).

Ptychohrancus (Simpson) Oetmann, 1912, p. 305; Simpson, 1914, p. 332; Elliy-
saria Feieeson, 1914a, p. 7.

Type Obliquaria fasciolaris Rafinesque.

Only the type-species known from Pennsylvania.

208

MEMOIRS OF THE CARNEGIE MUSEUM.

Ellipsaria fasciolaris (Rafinesque) (1820).

Ptychohranchus phaseolus (Hildreth) Simpson, 1941, p. 333; Ellipsaria fasciolaris

(Rafinesque) Frierson, 1914a, p. 7; Ptychohranchus fasciolaris (Rafinesque)

Vanatta, 1915, p. 554.

Plate XIII, figs. 1, 2, 3.

Records fro7n Pennsylvania:

Harn, 1891 (western Pennsylvania).

Marshall, 1895 (Allegheny River, Warren Co.).

Rhoads, 1899 (Beaver River, Wampum, Lawrence Co.).

Ortmann, 19096, pp. 194 and 202.

Characters of the shell: Shell of medium size, but comparatively thick and
heavy. Outline subelliptical, more or less elongated; when old, often humped
(subtriangular) , and drawn out at the lower posterior end. Upper and lower
margins more or less convex, posterior end narrower than anterior, rounded or
bluntly pointed. In old specimens the lower margin may be straight, and it
sometimes becomes even concave in the middle. Beaks in front of the middle,
very little elevated above the hinge-line. Beak-sculpture rudimentary, consisting
of two or three faint ridges which are interrupted, and have thus a tendency to
appear double-looped.

Valves moderately and rather evenly convex, slightly more flattened on the
sides. Posterior ridge practically absent, or indicated only by a stronger con-
vexity of the valves. No sculpture upon the surface.

Epidermis yellowish to light brown, rarely dark brown; generally ornamented
with dark green rays, which are most distinct in young specimens. The rays are
capillary and interrupted, but are generally grouped together in bundles, and
very often fused together, so as to form more or less squarish, green spots. There
is great variability in the size and the arrangement of these spots, and in very
rare cases they are entirely missing. Old shells have the rays less distinct, and
the epidermis becomes more or less uniformly brown. Concentric bands of color
are absent, or the color is only slightly darker along the growth-lines.

Hinge well-developed. Pseudocardinals stumpy, heavy, but not much ele-
vated. Laterals thick and heavy, club-shaped, thicker posteriorly. Interdentum
narrow. Beak-cavity shallow. Dorsal muscle-scars in the beak-cavity. Ad-
ductor-scars distinct and rather deep.

Nacre white, generally without any color, except in very young specimens,
where there may be a faint pink blush.

Externally there are no sexual differences in the shell, except that the males

OKTMANN: monograph of the naiades of PENNSYLVANIA.

209

are slightly more compressed than the females. It is indeed often impossible to
ascertain the sex by this character. Internally the female shell is marked by a
wide, oblique groove, running from the beak-cavity toward the posterior end, which
corresponds to the marsupium (PL XIII, fig. 2). The marsupial folds are also
generally marked as depressions in this groove. These depressions are best marked
in old specimens and are indistinct or absent in young ones.

L. H. D.

Size: 1. Cannelton, Cat. No. 61.1166 (9 ) 132 mm. 72 mm. 43 mm.

2. Harbor Bridge, Cat. No. 61.3601 (gravid 9 ) . . . . 125 “ 62 “ 40 “

3. New Sheffield, Cat. No. 61.3275 (cT) 109 “ 65 “ 32 “

4. Shenango, Cat. No. 61.4134 (c?) 87 “ 47 “ 28 “

5. Rose Point, Cat. No. 61.3116 (d") 73 “ 42 “ 22 “

6. Waterford, Cat. No. 61.4133 ( 9 gravid) 66 “ 36 “ 21 “

Our largest specimens (Nos. 1 and 2) exceed the maximum size (110 mm.)
recorded by Scammon (1906, p. 320).

Soft parts (See Ortmann, 1912, p. 306). They have been figured by Lea (Obs.
VII, 1860, PL 29, fig. 101) and Lefevre & Curtis (1910, PL 1, fig. 1, and 1912, PL 6,
fig. 1). Glochidia: Lea (Obs. VI, 1858, PL 5, fig. 12) poor; Ortmann (19115, PL
89, fig. 14).

Breeding season: The following records for gravid females are at hand: Aug.
4, 1908 (eggs); Aug. 29, 1910; Sept. 1, 1908 (young glochidia); Sept. 2, 1907;

Sept. 6, 1908; Sept. 7, 1908; Sept. 11, 1913; Sept. 13, 1909; Sept. 13, 1915;

Sept. 13, 1917; Sept. 15, 1908; Sept. 15, 1915; Sept. 18, 1917; Sept. 21, 1907;

Sept. 21, 1908; Sept. 27, 1909; Oct. 4, 1910; Oct. 15, 1907; Oct. 15, 1908; Oct.

19, 1908; Oct. 21, 1908; Oct. 23, 1907. Then again: June 23, 1910 (discharging);
Aug. 31, 1906 (discharging).

The beginning of the season is well established as being the month of August,
and the discharge takes place in spring and early summer. Probably the (single)
discharging individual observed on Aug. 31 was exceptionally belated. The
species is positively bradytictic.

Remarks: This is an easily recognized species. The subelliptical outline,
light brown epidermis, color-markings, and heavy hinge are unmistakable. How-
ever specimens of dark color and without rays might be mistaken for Elliptio
dilatatus, but the interior of the shell generally determines the question.

There is a good deal of variation in shape and color. The humped shape of
old shells does not develop in many cases, although on the other hand it may be
already in evidence in comparatively small individuals. Specimens from the
mountain-streams (Cheat, Loyalhanna, Quemahoning) are much smaller than

210

MEMOIRS OF THE CARNEGIE MUSEUM.

usual, and thus there is in the mountains a tendency toward the formation of a
small race. Lake Erie possesses a somewhat peculiar form (Plate XIII, fig. 3),
in which the specimens remain small; the dimensions of the largest at hand, a
gravid female, being: L. 72, H. 40, D. 24 mm., and they assume the humped shape
when comparatively small. Further, the gTOwth-lines are more crowded, and
more regular, but not very distinct (not marked by color). Young shells of the

■ Ellipsaria fasciolaris.

• Obliquaria reflexa.

+ Cyprogenia stegaria.

X Do. (Indian garbage heap).

form from Lake Erie are indistinguishable from specimens of the same size from
the Ohio-drainage. I do not think it advisable to distinguish the lake-form by a
varietal name, but the tendency to develope a local race is clearly indicated in
this case.

Specimens from Conneaut Creek (tributary to Lake Erie) are entirely normal.
It should be mentioned that the form from Chautauqua Lake in New York
inclines toward that of Lake Erie in the more crowded growth-lines, and the average
size, a fact which is apparently due to the parallel influence of the environment.

Localities in Pennsylvania, represented in the Carnegie Museum:

Small tributaries of Ohio:

Little Beaver Creek, Cannelton (Miss Vera White & H. H. Smith) and New Galilee, Beaver Co.; Enon
Valley, Lawrence Co.

Raccoon Creek, New Sheffield, Beaver Co,

ORTMANN; monograph of the naiades of PENNSYLVANIA.

211

Beaver-drainage :

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Ellwood City, Lawrence Co. (G. H. Clapp & H. H. Smith).

Slipperyrock Creek, Wurtemberg and Rose Point, Lawrence Co.

Mahoning River, Mahoningtown, Coverts, Edinburg, and Hillsville, Lawrence Co.

Neshannock Creek, Eastbrook and Volant, Lawrence Co.; Leesburg, Mercer Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Sharpsville, Clarksville, Shenango and
Jamestown, Mercer Co.

Pymatuning Cre.ek, Pymatuning Township, Mercer Co.

Little Shenango River, Greenville, Mercer Co.

Allegheny-drainage :

Allegheny River, Kelly, Armstrong Co.; Walnut Bend, Venango Co.; Tionesta and Hickory, Forest Co.
Buffalo Creek, Harbison, Butler Co.

Loyalhanna River, Idlepark and Ligonier, Westmoreland Co.^^°

Quemahoning Creek, Stantons Mill, Somerset Co.

Little Mahoning Creek, Goodville, Indiana Co.

Sandy Creek, Sandylake, Mercer Co.

French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co.
Leboeuf Creek, Waterford, Erie Co.

Connewango Creek, Russell, Warren Co.

Monongahela-drainage:

Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).

Dunkard Creek, Wiley and Mount Morris, Greene Co.

Cheat River, Cheat Haven, Fayette Co.

Lake Erie-drainage:

Conneaut Creek, West Springfield, Erie Co.

Lake Erie, Presque Isle Bay, Erie, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Lake Erie, Vermilion, Erie Co., Ohio; La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).

Grand River, Cayuga, Haldimand Co., Ontario, Canada (C. Goodrich).

Sandusky River, Fremont, Sandusky Co., Ohio (C. Goodrich).

Maumee River, Roche de Boeuf Rapids and Waterville, Lucas Co.; Otsego Rapids, Wood Co.; Defiance,
Defiance Co., Ohio (C. Goodrich).

Raisin River, Monroe and Grape P. 0., Monroe Co., Michigan (C. Goodrich).

Ohio-drainage :

Lake Chautauqua, Bemus Point (D. R. Sumstine), Griffith Landing (Miss. B. Ortmann), and Celoron
(P. E. Nordgren), Chautauqua Co., New York.

Tuscarawas River, Ohio (Holland collection).

Ohio River, Toronto, Jefferson Co.; Portsmouth, Scioto Co., Ohio.

Dead shells were seen, but not taken, in the Conemaugh River at New Florence, Westmore-
land Co.

212

MEMOIRS OF THE CARNEGIE MUSEUM.

Cheat River, Jaco and Mont Chateau, Monongalia Co., West Virginia.

West Fork River, Lynch Mines, Harrison Co.; West Milford, Harrison Co. (W. F. Graham); Lightburn,
Lewis Co., West Virginia.

Little Kanawha River, Grantsville, Calhoun Co. (W. F. Graham); Burnsville, Braxton Co., West Virginia.
Elk River, Shelton and Clay, Clay Co.; Gassaway and Sutton, Braxton Co., West Virginia.

Licking River, Farmer, Rowan Co., Kentucky.

Tennessee-drainage :

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith); Knox Co., Tennessee (Smith
collection).

Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).

Elk River, Estill Springs, Franklin Co., Tennessee (H. H. Smith).

Paint Rock River, Paint Rock and Trenton, Jackson Co., Alabama (H. H. Smith).

South Chickamauga Creek, Ringgold, Catoosa Co., Georgia.

French Broad River, Boyd Creek, Sevier Co., Tennessee.

Holston River, Mascot, Knox Co.; Turley Mill, Noeton, and Holston Station, Grainger Co.; Austin
Mill, Hawkins Co., Tennessee.

South Fork Holston River, Pactolus, Bluff City, and Emmett, Sullivan Co., Tennessee.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., and Mendota,
Washington Co., Virginia.

Clinch River, Edgemoor, 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.; Fink and Cleveland, Russell Co., Virginia.

Emory River, Harriman, Roane Co., Tennessee.

Powell River, Combs, Claiborne Co., Tennessee.

Distribution and Ecology in Pennsylvania (See fig. 22) : In Pennsylvania this
is preeminently a species of smaller rivers and creeks. It is very rare in the Alle-
gheny below Oil City, and has not been found in the Ohio. In the Monongahela
it must also have been rare. But in the tributaries it is abundant, with the ex-
ception of the smallest headwaters. It goes far eastwards toward the divide in
Indiana, Westmoreland, and Somerset Counties. It is also found in Cheat River
and the upper Monongahela in West Virginia.

Furthermore it occurs in Lake Erie and its tributary, Conneaut Creek, and
may have crossed into the lake-drainage from the upper Beaver-drainage.

That it avoids large rivers holds good further down the Ohio, although it is
not entirely absent there. Its ecological preferences are distinctly for riffles and
strongly flowing water, with finer or coarser, but firmly packed gravel. Very
frequently it is found along the edges of Dfani/iera-patches, always in a lively
current. In Lake Erie it is chiefly found on the North shore of Presque Isle Bay,
in two or three feet of water, on sandy or slightly gravelly bottom, exposed to the
moderate surf of the bay, but it is not abundant there. Scammon (1906) says
that this species prefers (in Kansas) the smaller streams.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

213

General distribution: Type locality, Kentucky River (Vanatta) (Rafinesque
gives as habitat the Ohio, Wabash, and Kentucky Rivers).

Simpson (1900) gives its distribution as follows: “Ohio, Tennessee, and
Cumberland river systems; peninsula of Michigan; Kansas; Arkansas; Indian
Territory; Louisiana.” This possibly requires modification with regard to the
southwestern range. The most noteworthy feature in its distribution is its absence
in the upper Mississippi-drainage. Numerous localities are known from all over
Ohio (Sterki, 1907a). In Indiana (Call, 1896a & 1900) it is found practically all
over the state, both in the Ohio and Erie drainages, but it is absent in the Lake
Michigan-drainage. As our localities show, it occurs also in West Virginia, and
is very abundant in the upper Tennessee region. Wilson & Clark (1914) report
it from the upper Cumberland River. In very strong contrast to the foregoing
is its scarcity in Illinois. In the latter state it is known only in the southern parts
and in the Wabash River (Baker, 1906). Simpson extends the range in a south-
westerly direction across the Mississippi. (See also Call, 1895, for Arkansas, and
Scammon, 1906, for Kansas.) However, the specimens I have seen from Missouri,
Arkansas, and Oklahoma are not typical E. fasciolaris, and represent, as has been
recognized by Utterback (1916, p. 128) and hinted at by Wheeler (1918, p. 120), a
different species. This species is distinct in the rays, which are fine and capillary,
and never form blotches.

In a northeasterly direction this species has crossed over into the lake-drainage,
going probably by way of the Maumee-route. It has spread over practically the
whole lower peninsula of Michigan (as far north as Cheboygan Co.) (See Walker,
1892 & 1898). In Lake Erie it has reached New York at Buffalo (Marshall,
1895). It is clear that the other locality in New York, Chautauqua Lake, was
reached from the upper Allegheny.

Two records from Wisconsin and Minnesota are undoubtedly in error and
already have been dropped by Simpson.

Genus Obliquaria Rafinesque (1820).

Ortmann, 1912, p. 309; Simpson, 1914, p. 329.

Type Obliquaria refiexa Rafinesque.

A monotypic genus.

This species should be called ElUpsaria occidentalis (Conrad) (1836) (Currant River, Arkansas),
the synonym of which is Ptychobranchus cUntonensis Simpson (1900) (Archies Fork of the Little Red
River, Clinton, Arkansas). Unio occidentalis of Conrad has been entirely misunderstood by Simpson
(1914, p. 112, as Lampsilis occidentalis). The Carnegie Museum possesses material of this form from
Missouri, Arkansas, and Oklahoma. Its metropolis is in the Ozark Mountains.

214

MEMOIRS OF THE CARNEGIE MUSEUM.

Obliquaria reflexa Rafinesque (1820).

Obliquaria reflexa Rafinesque, Simpson, 1914, p. 330.

Plate XIII, fig. 4.

Records fro7n Pennsylvania:

Clapp, 1895 (Allegheny Co.).

Rhoads, 1899 (Oliio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.).

Ortmann, 19096, p. 193.

Characters of the shell: Shell rather small, hardly of medium size, thick and
solid. Outline more or less rounded, subovate or subtraiiezoidal, short and high,
more or less pointed at the lower posterior end. Anterior and lower margins
regularly rounded; lower margin ascending posteriorly, straight or slightly con-
cave. Upper margin short, passing in a curve or blunt angle into the obliquely
descending posterior margin. Beaks near, but somewhat in front of the middle,
elevated over the hinge-line and incurved. Beak-sculpture consisting of a few
(two to three) rather heavy, l)ut not sharply defined, concentric bars, which have
an indistinct tubercle upon the posterior ridge. Valves flat or convex, with a
more or less distinct, rounded posterior ridge, generally with a shallow radial
furrow,, which may be obsolete. In front of this furrow stands a row of large,
lirominent knobs, which alternate with each other on the ojiposite valves. They
are rounded, conical, or vertically compressed. Maximum number of these knobs
on each valve, four or five. Posterior slope often ornamented with short, corru-
gated ridges.

Epidermis yellowish to brown, with indistinct darker concentric bands.
There may be rays, fine, wavy, or broader; they sometimes spread over the surface,
so that the whole epidermis appears dark green. In old specimens the epidermis
is often uniformly brown.

Hinge well-developed. Pseudocardinals large, ragged, triangular, two in the
left, one in the right valve. Laterals thick and short. Interdentum short and
rather narrow. Beak cavity moderate. Dorsal muscle-scars in the beak-cavity.
Anterior adductor-scars small, deeply impressed, posterior opes also small and
distinct, but less impressed. Nacre silvery white, very rarely colored: I have
seen only a few individuals from Alabama with purple nacre.

Sexual differences in the shell very uncertain. On the average, the females
are more swollen, with the radial furrow indistinct, and the posterior lower margin
not emarginate; while the male shells are more compressed, with the furrow more
distinct, and the emargination of the lower margin more frequently developed.
These differences, however, are slight, and there are shells in which the sex cannot

ORTMANN: monograph of the naiades of PENNSYLVANIA,

215

positively be determined by these characters. The females seem to remain smaller

than the males.

L. H. D.

Size: 1. Industry, Cat. No. 61.4423 (cf) 63 mm. 53 mm. 33 mm.

2. Cooks Ferry, Cat. No. 61.3583 (d") 56 “ 47 “ 28 “

3. Shippingport, Cat. No. 61.4756 (cf) 56 “ 48 “ 27 “

4. Industry, Cat. No. 61.4122 (cf) 46 “ 38 “ 22 “

5. Industry, Cat. No. 61.3585 (9 ) 39 " 33 “ 20 “

No. 1 is the largest at hand, surpassing the maximum length given by Scam-
mon (1906).

Soft parts figured by Lefevre & Curtis, 1910, PL 1, fig. 3; 1912, PL 7, fig. 7;
described and figured by Ortmann, 1912, p. 310, fig. 16. Glochidia figured by
Lefevre & Curtis, 1910, p. 97, fig. M; 1912, p. 146, fig. M; Ortmann, 1912, PL 20,
fig. 1; Surber, 1912, PL 2, fig. 39. According to Lefevre & Curtis the measure-
ments are: 0.225 X 0.230; Surber: 0.225 X 0.235; Ortmann: 0.22 X 0.22 mm.

Breeding season: According to Lefevre & Curtis (1912), and also Surber
(1912), the breeding season lasts from May to August: embryos are present from
the end of May to July 9, and glochidia from June 20 to Aug. 8. Thus this species
would appear to be tachytictic.

I have specimens with glochidia (and discharging), collected on May 19,-1911;
June 20, 21, 22, 1911; July 13, 1911; July 29, 1914; and Aug. 6, 1910. These
accord with the breeding season as given above, with the exception of the first
date (May 19). This specimen is from Arkansas. It may be an exceptionally
early date, or the breeding season may begin earlier in the South.

It would be well to try to obtain additional data. The species is adapted to
a long breeding season. Being a rather primitive form, we may have here primitive
or transitional conditions; or on the other hand the breeding season may have
been re-adapted to more southern conditions, and may have become irregular.

Remarks: A very peculiar, ancient type in the subfamily, which cannot be
mistaken for any other species on account of the peculiar shape and the unique
sculpture. It is rather variable in outline and color.

There is a dwarf race in Lake Erie, light in color, which possibly deserves a
varietal name; but since this form has not as yet been found in Pennsylvania, I
shall not treat of it here.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Shippingport, Cooks Ferry, and Industry, Beaver Co.

Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).

216

MEMOIRS OF THE CARNEGIE MUSEUM.

Other localities represented in the Carnegie Museum:

Lake-drainage :

Lake Erie, Maumee Bay, Toledo, Lucas Co., Ohio; and La Plaisance Bay, Monroe Co., Michigan (C.
Goodrich).

Ohio-drainage:

Ohio River, Toronto, Jefferson Co., Ohio; St. Marys, Pleasants Co., West Virginia; Portland, Meigs
Co., Ohio; Portsmouth, Scioto Co., Ohio.

Pocatalico River, Raymond City, Putnam Co., West Virginia.

Tennessee-drainage :

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).

Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).

Clinch River, Solway, Knox Co.; Edgemoor and Clinton, Anderson Co., Tennessee.

Western range:

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection); Moline, Rock Island Co.,
Illinois (P. E. Nordgren).

Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).

Spring River, Black Rock, Lawrence Co., Arkansas (A. A. Hinkle}^-
White River, Cotter, Baxter Co., Arkansas (A. A. Hinkley).

Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).

Verdigris River, Inola, Rogers Co., Oklahoma (F. B. Isely).

Bayou Pierre, De Soto Parish, Louisiana (L. S. Frierson).

Sabine River, Logansport, De Soto Parish, Louisiana (L. S. Frierson).

Alabama-drainage:

Forks of Black Warrior River, Walker Co., Alabama (H. H. Smith).

Coosa River, Wetumpka, Elmore Co.; Weduska Shoals and Wilsonville, Shelby Co.; Coosa Valley,
Riverside, and Lock 4, St. Clair Co.; Fomby Shoals, Calhoun Co.; Greensport, St. Clair Co.;
Minnesota Bend, Cherokee Co., Alabama (H. H. Smith).

Distribution and Ecology (See fig. 22) : Type locality, Ohio River, Letart Falls,
Meigs Co., Ohio (Rafinesque) .

In Pennsylvania this species is restricted to the large rivers, the Ohio and
Monongahela. I found it repeatedly in Beaver County, but it is rather rare.
Farther down it becomes more abundant in the Ohio. In general, it seems to be
restricted to the Ohio proper and a few of its larger tributaries, as the Muskingum
(Dewey, 1856) and Scioto. Sterki (1907a) says that it is not found in the Tus-
carawas, but in the Mahoning River. However I doubt the latter record, since
Dean (1890) does not mention it, and since it is absent in the Beaver and Mahoning
in Pennsylvania. The smallest stream in which I found this species, is the Poca-
talico River in West Virginia, tributary to the Kanawha, and here it was scarce,
and only occurred in the lower part of the stream. In Indiana also this species

ORTMANN: monograph of the naiades of PENNSYLVANIA.

217

is chiefly found in the larger rivers (Ohio, White, Wabash, Kankakee, according to
Call, 1896a & 1900). It is more abundant in Illinois (Baker, 1906; Forbes &
Richardson, 1913), and here it crosses over into the drainage of Lake Michigan
(Calumet River, Baker, 1898a). It goes up the Mississippi in Illinois and Iowa,
and reaches Wisconsin in the Fox River (Barnes, 1823), and southern Minnesota
(Grant, 1886; Holzinger, 1888).

Southward it is found in the western tributaries of the Mississippi in eastern
Kansas (Scammon, 1906), Arkansas (Call, 1895; Wheeler, 1918), Oklahoma and
Louisiana (Vaughan, 1893; Frierson, 1899) and also in eastern Texas (Singley,
1893).

In the southern tributaries of the Ohio it has been reported from the Ken-
tucky River (Rafinesque) . It is common in the Cumberland (Wilson & Clark,
1914) ; and, in the Tennessee, it ascends in the region of Knoxville into the lower
Clinch River (Carnegie Museum).

It turns up in the Alabama-drainage (Lewis, 1870 and 1877; Call, 1885;
Carnegie Museum) , and there goes as far as the Etowah River in northern Georgia
(Call, 1885).

In the north it has been reported in Michigan from the Grand and Saginaw
Rivers (Walker, 1894 & 1898), and it is also found in Lake Erie in Michigan and
Ohio (Sterki, 1907a, and Walker, 1913). It has not been found in Pennsylvania,
and Calks record (1895, p. 12) from western New York has not been substantiated
by positive information. (It does not occur in the list of Marshall, 1895.)

It is clear that the species has crossed over into the lake-drainage from the
upper Ohio system; but how it reached the Alabama-drainage rema’ns to be
investigated.

It is evident that it prefers large rivers, and it reaches Pennsylvania only
in the large streams. According to my observations, it is found chiefly in the
deep channel of the Ohio, upon the shell-banks, and is regularly taken by the
clam-diggers. Scammon (1906) gives gravel-beds as the favorite habitat. But
it seems to be also found on muddy bottoms (Baker, 1898a), and Call (1900) says,
that it is almost ubiquitous with regard to its habitat. In Pocatalico River I
found it in pure, shifting sand.

Genus Cyprogenia Agassiz (1852).

Ortmann, 1912, p. 212; Simpson, 1914, p. 326.

Type Obovaria stegaria Rafinesque.

Only one species known from Pennsylvania.

218

MEMOIKS OF THE CARNEGIE MUSEUM.

Cyprogenia stegaria (Rafinesque) (1820).

Cy'progenia irromta (Lea) Simpson, 1914, p. 326; Cyprogenia stegaria (Rafinesque)

Vanatta, 1915, p. 554.

Plate XIII, fig. 5.

Records fro7n Pennsylvania:

Rhoads, 1899 (Oliio River, Beaver, Beaver Co.)

Ortmanii, 19096, p. 193.

Characters of the shell: Shell hardly of medium size, but solid. Outline sub-
circular, subtrapezoidal, or rounded triangular. Anterior and lower margins more
or less rounded, but the latter tending in the posterior part to be straight or slightly
concave. Ujiper margin short, slightly convex, forming a more or less distinct
angle with the posterior margin, which is nearly vertical. Beaks slightly anterior
to the middle, moderately inflated and elevated. Beak-sculpture rudimentary,
consisting of a few slightly double-looped bars. Valves more or less convex,
sometimes making the shell subglobular. Young specimens have a distinct, but
rounded, posterior ridge, which liecomes effaced in old ones. Generally there is a
slight radial groove in front of the ridge, almost effaced in old specimens. Surface
adorned by flat nodules, most distinct upon the posterior ridge and in front of the
groove. Anterior part of the shell smooth, but there may be nodules in the groove
and upon the posterior slope, where they may assume a radial arrangement. In
old shells the nodules are much obliterated. The growth-rests are often marked
by concentric, low ridges.

Epidermis light green or yellow, to light brown, ornamented with green
mottlings, which fall into finer or broader, interrupted rays. The broad rays
prevail upon the anterior part of the shell, the fine ones upon the posterior. In
old shells the mottlings become indistinct. Concentric color-markings are absent
or very faint.

Hinge well-developed. Pseudocardinals heayjq triangular, blunt and ragged,
two in left, one in the right valve. Interdentum ver\^ broad and short. Laterals
rather short, heavAG Beak-cavity moderately deep. Dorsal muscle-scars in the
beak-cavity, upon the hinge-plate. Adductor-scars distinct, small, deeply im-
pressed. Nacre silvery white, very rarely with a faint blush of pink.

I cannot see any sexual differences in the shell.

L. H. D.

Size: I. Godfrey, Cat. No. 61.4760 62 mm. 57 mm. 38 mm.

2. do. Cat. No. 61.3581 56 “ 54 “ 34 "

3. Industry, Cat. No. 61.4422 54 “ 52 “ 33 “

I have not seen any larger specimens than No. 1.

4

ORTMANN: monograph of the naiades of PENNSYLVANIA. 219

The soft parts were described and figured by Lea (Obs. I, 1834, PL 5, figs. 6, 7),
but fig. 7 is wrong; also (Obs. X, 1863, p. 433). Ortmann (1912, p. 313, fig. 17),
and Lefevre & Curtis (1912, PL 7, fig. 8). The GlocMdia were described by Sterki
(1898, p. 19) and figured by Ortmann (1912, PL 19, fig. 6) and also by Surber
(1912, PL 1, fig. 11). They measure, according to Sterki, 0.21 X 0.17; according
to Surber, 0.210 X 0.185; according to Ortmann, 0.18 X 0.15 mm.

Breeding season: The species is hradytictic according to Lefevre & Curtis
(glochidia in November). Surber found gravid females in October and November.
I found them with eggs on Sept. 7 and 12, 1914, and Sept. 14, 1915, and Sept. 17,
1915, and Sept. 24, 1915.

Remarks: A well-marked species, distinguished by its somewhat subglob ular
shape, nodular surface, and the peculiar, mottled character of the color of the
epidermis. The shape, however, is rather variable, and also the development of
the nodules. The placentae are mostly red, but I have found specimens (in Clinch
River) with white placenta.

Localities represented in the Carnegie Museuyn:

Ohio River, Industry, Beaver Co., Pennsylvania.

Allegheny River, Natrona, Allegheny Co.; Aladdin and Godfrey, Armstrong Co., Pennsylvania.
Tuscarawas River, Ohio (Holland collection).

Little Miami River, Xenia, Greene Co., Ohio (C. Goodrich).

Ohio River, St. Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland,
Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio; and Cincinnati, Hamilton Co., Ohio (Juny col-
lection).

Cumberland River, Cloyds Landing and Albany Landing, Cumberland Co., Kentucky (B. Walker, donor).
Tennessee River, Tuscumbia, Colbert Co., and Florence, Lauderdale Co., Alabama (H. H.- Smith);

Knoxville, Knox Co., Tennessee (Hartman collection).

Holston River, Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill, Grainger Co., Tennessee.

Clinch River, Solway, Knox Co.; Edgemoor, Clinton, and Offutt, Anderson Co.; Black Fox Ford, Union
Co.; Clinch River Station, Claiborne Co., Tennessee.

Distribution and Ecology (See fig. 22): Type locality, Ohio River (Rafinesque) .
In Pennsylvania this species has been found in the Ohio and lower Allegheny.
All specimens collected by myself (only seven) were dead shells, but some of
them were quite fresh. However, this species must once have also existed in the
Monongahela, at least as far up as the mouth of Cheat River, for I have found
specimens in an old Indian garbage heap,^^^ opposite Point Marion (See Ortmann,
1909c, p. 13).

According to Simpson (1900) it belongs to the Ohio, Cumberland, and Tennes-
see Rivers, and is mainly restricted to these large rivers, and a few of their larger

Kitchen-midden. -Editor.

220

MEMOIRS OF THE CARNEGIE MUSEUM.

tributaries. Thus, for instance, it occurs in Ohio, in the Muskingum at Marietta
(Hildreth, 1828), in the Tuscarawas (Dean, 1890; Sterki, 1907a), and in the
Scioto and Great Miami (Sterki). Sterki reports it also from the Mahoning
River, but it has never been found in the Pennsylvanian part of it, and never
anywhere else in the Beaver-drainage. In Indiana it is found besides the Ohio, in
the White and Wabash Rivers (Call, 1896a and 1900). In Illinois it occurs only
in the Wabash (Baker, 1906). A number of localities are known in Kentucky,
Tennessee, and northern Alabama in the Cumberland and Tennessee systems
(See Wilson & Clark, 1914).

West of the Alississippi it has been reported only by Call (1895) from the St.
Francis River, Wittsburg, Cross Co., and Saline River, Benton, Saline Co., Ar-
kansas, but this is questioned by Simpson, and correctly, as I think. I know that
this species is represented in this region (Missouri, Kansas, Arkansas, Oklahoma)
by another, C. aberti (Conrad), which has apparently been mistaken for the one
under consideration.

In the Ohio between Pittsburgh and Cincinnati this species is found in the
mussel-beds in the deep channel of the river, on gravelly bottoms with steady
currents. In the Tennessee-drainage I found it frecpiently in firmly i)acked gravel,
in strongly flowing water, in rivers of medium size (Clinch, Holston).

Genus Obovaria Rafinesque (1820).^^^

Ortmann, 1912, p. 320; Simpson, 1914, p. 289.

Type Unio retusa Lamarck.

The genus is divided into two subgenera.

Key to the Shbgenera of Obovaria.

Ui. Shell rounded or subovate, upright, with nearly central, or anteriorly incurved beaks. Pseudo-
cardinals divergent, not parallel to the laterals. Epidermis more or less brown. Nacre often red

or purplish Subgenus Obovaria.

az. Shell ovate or elliptical, oblique; beaks quite anterior. Pseudocardinals in adult specimens sub-
parallel to the laterals. Epidermis greenish. Nacre white Subgenus Pseudoon.

Subgenus Obovaria Simpson (1900).

Ortmann, 1912, ji. 321; Simpson, 1914, p. 290.

Type Obovaria {Unio) retusa Lamarck.

Two species and one variety are found in Pennsylvania.

Key to the Forms of Obovaria.

tti. Shell as high as, or higher than, long. Beaks much elevated and much incurved. Nacre deep purple
with white margin 0. retusa.

Not “ 1819,” as Simpson gives the date. In 1819 Rafinesque published this name, but as “ nomen
nudum.”

ORTMANN: monograph of the naiades op PENNSYLVANIA.

221

a 2- Shell not higher than long. Beaks less elevated and not much incruved. Nacre white or purplish.

bi. Shell swollen. Diameter sixty percent of the length or more 0. suhrotunda.

hi. Shell compressed. Diameter less than sixty percent of length 0. subrotunda levigata.

Obovaria (Obovaria) retusa (Lamarck) (1819).

Obovaria retusa (Lamarck) Simpson, 1914, p. 290.

Plate XIII, figs. 6, 7.

Records from Pennsylvania:

Ortmann, 19096, p. 192.

Characters of shell: Shell of medium size, heavy and solid. Outline subcircular
or subovate, about as high as long, or even higher than long, upright. Anterior,
lower, and posterior margins almost regularly rounded, only with a blunt lower
posterior angle in the male, and a slight emargination in this region in the female.
Upper margin short, convex. Beaks swollen and elevated, situated near the middle
of the shell, but strongly incurved forward. Beak-sculpture rudimentary, never
distinctly seen by the writer. Valves strongly convex, without a distinct posterior
ridge. No sculpture on the outer surface, but irregular concentric ridges, corre-
sponding to the growth-lines, are often present.

Epidermis light to dark brown, generally quite uniformly dark brown upon
the disk, while the posterior slope is lighter, pale brown or yellowish. The lighter
color is not sharply marked off from the darker part, and often the contrast is
not evident at all. Upon the lighter posterior slope, there are sometimes a few
feeble greenish rays, and in young specimens, such rays may be indicated by mere
traces upon the disk, but normally there are no rays at all.

Hinge well-developed. Pseudocardinals very heavy, stumpy, ragged, tri-
angular, one in right, two in left valve, the latter divergent and not parallel to
the laterals. Interdentum broad and short. Laterals short, thick and heavy.
Beak-cavity rather deep and narrow. Dorsal muscle-scars upon the inside of the
hinge-plate. Adductor-scars distinct and deeply impressed. Nacre of various
shades of purple, generally very deeply colored, but the margin outside of the
mantle-scar is generally white.

Sexual differences of the shell moderate, but distinctly marked. In the
male, the lower and posterior margins form a regular curve or meet in an indistinct,
rounded lower posterior angle. In the female the disk shows a slight swelling,
forming a slight projection in the posterior part of the lower margin situated at a
lower level than the posterior angle of the male. Above this swelling, and behind
it, the shell is slightly depressed, and the posterior margin ascends straight or is

222

MEMOIES OF THE CAKNEGIE MUSEUM.

somewhat concave, before it reaches the level of the posterior angle of the male.
Thus the female shell has two blunt posterior angles on the hind end, but this
structure may be rather obscure. The female shell is generally higher than that
of the male, and the male seems to grow to a slightly larger size than the female.

L. H. D.

Size: 1. Industry, Cat. No. 61.3556 (cf) 58 mm. 58 mm. 37 mm.

2. do. “ “ 61.3555 (gravid 9 ) 58 “ 64 “ 39 “

Soft 'parts (See Ortmann, 1912, p. 321, 322, fig. 20). Glochidia: Ortmann,
1912, PL 19, fig. 9; Surber, 1912, PI. 3, fig. 47. Measurements: 0.22 X 0.27
(Ortmann); 0.240 X 0.295 (Surber).

Breeding season: On Aug. 29, 1908 and Sept. 7, 1914, I found specimens with
eggs, and on Sept. 22, 1910 specimens with glochidia. This indicates the beginning

• Ohovaria retusa.

m Ohovaria olivaria. '

+ Ohovaria suhrotunda.

X Ohovaria suhrotunda levigata.

> Ohovaria suhrotunda levigata (from Indian garbage heap).

of the season in autumn. Surber (1912) reports this species as gravid in September.
It probably is hrad'ytictic.

Remarks: There is no trouble in recognizing this species; the shape and the
color of the nacre are quite unique.

Localities represented in the Carnegie Museum:

Oliio River, Industry, Beaver Co., Pennsylvania.

Ohio River, Toronto, Jefferson Co., Oliio; Wheeling, Ohio Co., West Virginia; (W. F. Graham); St.
Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland, Meigs
Co., Ohio; Portsmouth, Scioto Co., Oliio.

Tennessee River, Knox Co., Tennessee (Smith collection).

Clinch River, Clinton, Anderson Co., Tennessee.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

223

Distribution and Ecology (See fig. 23) : Type locality, unknown. The original
type locality (Nova Scotia) is erroneous.

Only two specimens have ever been found in Pennsylvania, both at the same
place in Beaver Co. This marks the farthest point upstream in the Ohio of the
range of this species.

Simpson (1900) gives the range as “Ohio, Cumberland, and Tennessee River
systems.” To these it seems to be restricted, and in the middle and upper Ohio,
it is also restricted to this river, and does not go into the tributaries (For Ohio see
Sterki, 1907a). In Indiana and Illinois, however, it has spread into the larger
affluents. Call (1896a, and 1900) reports it for Indiana from the Wabash, White,
Whitewater, Patoka, Eel, and Kankakee, and Baker (1906) gives it for Illinois
from the Wabash and Spoon Rivers, as far north as La Salle County. It is also
found in the lower Ohio (in Indiana and Illinois), and in the Mississippi River,
according to Call (1895, p. 47) “north of Arkansas,” but it is missing in Witter’s
list (1878) of the shells from Muscatine, Iowa, and in Pratt’s list (1876) from
Davenport, Iowa.

0. retusa is known from the Cumberland River (Wilson & Clark, 1914; also
previously reported from Nashville, Davidson Co., Tennessee, by Call, 1895).
In the Tennessee it goes up to the region of Knoxville, entering here the lower
Clinch.

It is unknown from west of the Mississippi and from the Gulf-drainage, and
thus the species is rather restricted in its distribution, and is apparently a form of
the larger rivers. In the Ohio between Pittsburgh and Cincinnati I found it asso-
ciated with the bank-forming shells, and it is here regularly taken by the clam-
diggers, but rejected on account of the color of the nacre. In a few cases, I col-
lected it in smaller branches of the Ohio in fine gravel, most abundantly at Portland,
Ohio.

Obovaria (Obovaria) subrotunda (Rafinesque) (1820).

Obovaria circulus (Lea) Simpson, 1914, p. 291; Obovaria subrotunda (Rafinesque)

Vanatta, 1915, p. 552.

Plate XIV, figs. 1, 2.

Records from Pennsylvania:

Lea, Obs. I, 1834 (Monongahela River, Pittsburgh).

Harn, 1891 (western Pennsylvania).

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co.) (reported as U. lens).

Ortmann, 1909&, p. 192.

Also from the Maumee-drainage (St. Joseph River) but this surely needs confirmation. It is
also said to be present in a few small lakes in northern Indiana.

224

MEMOIRS OF THE CARNEGIE MUSEUM.

Characters of the shell: Shell rather small, but thick and solid. Outline irre-
gularly subcircular, about as high as long, or slightly longer than high, upright.
Anterior and lower margins forming a regular curve. Lower margin ascending
posteriorly, more or less straight, meeting the obliquely descending posterior
margin in a blunt angle in the male. In the female this angle is less distinct, and
the posterior margin becomes nearly vertical. Upper margin short, convex.
Leaks more or less swollen, slightly elevated, situated close to the middle of the
shell, not incurved. Beak-sculpture rudimentary, consisting of about four or
five weak bars, slightly sinuated in the middle, angled behind, and disappearing
upon the posterior slope. Shell swollen, diameter at least sixty percent of the
length. Convexity of valves rather uniform, without any posterior ridge. No
sculpture upon the disk.

Epidermis light to dark brown, in young specimens sometimes yellowish
green; the disk is generally uniformly dark brown, while the posterior slope is
light brown or yellowish, sharply contrasted with the disk, but the contrast be-
comes less distinct toward the lower margin of the shell. In light-colored young
shells, there are sometimes faint traces of greenish rays, but generally rays are
entirely absent.

Hinge well-developed. Pseudocardinals strong, stumpy, ragged, triangular,
one in right, two in left valve, the latter divergent, and not parallel to the laterals.
Interdentum moderate or narrow. Laterals short, strong. Beak-cavity moder-
ately deep. Dorsal muscle-scars partly in the beak-cavity, partly upon the hinge-
plate. Adductor-scars distinct, and rather deeply impressed. Nacre silvery
white, or with more or less pink or light purple inside of the marginal zone.

Sexual differences in the shell present, but not strongly pronounced, and
sometimes obscure. In the male the posterior part of the lower margin ascends
and meets the posterior margin in a blunt angle, which, however, may be very
indistinct. In the female the lowermost point of the lower margin is situated
more backward, and then the margin ascends more suddenly, curving up into the
posterior margin more gradually, without forming a distinct angle. Thus the
posterior margin appears more vertical, the shell is rather truncate posteriorly,
and is rather more elevated. Besides, there is distinctly a difference in size, the
female being considerably smaller than the male.

L.

H.

D.

Pr.ct.

(Males) 1. Neville Island, Cat. No. 61.1798. .

. . .59

mm. 52

mm. 38 mm. .64

2. do. Cat. No. 61.1598. .

. . .49

“ 45

“ 32 “

.65

(Females) 3. Charleroi, Cat. No. 61.2840

. . .50

“ 48

« 34 «

.68

4. do. “ “ do

. . .38

“ 37

“ 24 “

.63

ORTMANN: monograph op the naiades of PENNSYLVANIA.

225

Soft parts (See Ortmann, 1912, p. 323). GlocMdia identical with those of var.
levigata, 0.20 X 0.23 mm., while Surber (1915, p. 7, fig. 8) gives: 0.170 X 0.215 mm.

Breeding season: 1 found the typical suhrotunda gravid with glochidia on
Sept. 22, 1910, and gravid and discharging on May 25, 1911. A female with eggs
was found on Sept. 6, 1914. Surber gives June 9, 1913 for glochidia. This would
agree with the bradytictic character of the species. A single individual, however,
discharging glochidia, was received from A. A. Hinkley, collected on Aug. 9, 1912
in the Wabash River. Probably this specimen was exceptionally belated.

Remarks: The more or less circular outline and subglobular shape, the almost
central beaks, and the light color of the posterior slope, serve to distinguish this
species. But there is great variability in the outline, and also in the convexity of
the valves, and, according to the latter, a variety {levigata) must be distinguished
from the main species, which will be treated below, and it should be pointed out,
that the latter is closely connected with the normal form by intergrades.

Another “species,” distinguished by Lea (and Simpson), 0. leihi (Lea), is
nothing but the form from Lake Erie of 0. subrotunda. It differs merely in its
smaller size (largest at hand: L. 42, H. 38, D. 26 mm.), and has, like many lake-
forms, more regular and more distinct growth-rests. It is also sometimes lighter
in color. I shall not treat of it here in detail, since I have not yet found it on the
Pennsylvanian shores of the lake.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Neville Island, Allegheny Co.

Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Other localities represented in the Carnegie Museum:

Ohio River, St. Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland,
Meigs Co., Oliio.

Wabash River, New Harmony, Posey Co., Indiana (A. A. Hinkley).

West Fork White River, Riverside, Greene Co., Indiana (J. D. Haseman).

Little Kanawha River, Grantsville, Calhoun Co., West Virginia (W. F. Graham).

Elk River, Shelton and Clay, Clay Co.; Gassaway and Sutton, Braxton Co., West Virginia.

Holston River, Mascot, Knox Co., Tennessee.

Distribution and Ecology (See fig. 23): Type locality, Ohio River (Rafinesque).
(Vanatta says: “Kentucky River”.)

In Pennsylvania the typical 0. subrotunda is restricted to the Ohio, the Beaver,
and the lower Monongahela, and it should be kept in mind, that just in this region
more compressed specimens turn up, which establish the transition toward the

226

MEMOIRS OF THE CARNEGIE MUSEUM.

var. levigata. Farther up only the latter is found, but farther down the Ohio,
0. subrotunda is common, and goes up into some of the tributaries (Little Kanawha
and Elk Rivers). The form from Elk River is peculiar, dwarfed, but with the
average diameter of sixty-one percent of the length. Sterki (1907a) believes
that circulus {subrotunda) is the male, and lens {levigata) the female. This is not
correct, for relying on Lea’s figures, just the opposite would be true. On account
of this confusion it is not possible to trace the exact distribution of the two forms
in Ohio.

In the rest of the range the two forms have not been kept strictly separate.
It suffices here to say, that the range of the true subrotunda covers, as Simpson
(1900) says: “the Ohio, Tennessee, and Cumberland river systems,” that it does
not extend west of the Mississippi (Call, 1885), and that it is restricted to larger
rivers. This is substantiated by the material in the Carnegie Museum from
West Virginia, Kentucky, and Tennessee. Wilson & Clark (1914) report this
form from the upper Cumberland.

0. subrotunda has also been reported from the south in the Alabama and
Tombigbee drainages (Lea and Lewis, 1870), but this is questioned by Simpson.
A few specimens from this region in the Carnegie Museum resemble 0. subrotunda
very much, but I have not enough material to decide the question.

This species also crosses over into the lake-drainage in southern Michigan
and Ohio. Specimens from Lake Erie proper form a peculiar race (var. leibi), and
appear more closely related to subrotunda, than to var. levigata. In the tributaries
of the lake (Maumee-drainage) the var. levigata is present. (See below.)

0. subrotunda in the Ohio prefers the shell-banks, in gravel and steady currents.
It is also found in small branches of the river, in the gravel of the riffles.

Obovaria (Obovaria) subrotunda levigata (Rafinesque) (1820).
Obovaria lens (Lea) Simpson, 1914, p. 293; Obovaria levigata (Rafinesque) Van-

ATTA, 1915, p. 552.

Plate XIV, figs. 3, 4.

Records from Pennsylvania :

Rhoads, 1899 (specimens from Beaver River, Wampum, Lawrence Co.).

Ortmami, 19096, p. 192 (included in circulus).

Characters of the variety: Shell similar to that of typical subrotunda, but more
compressed, the diameter less than sixty percent of the length. Sometimes, but

But in Elk River also the diameter decreases in the upstream direction. At Sutton specimens
were found, which should be called var. levigata according to the diameter.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

227

not always, the shell is of a lighter color. When lighter, the color is greenish or
brownish olive upon the disk, and yellowish upon the posterior slope.

L. H. D. Pr.ct.

Size: (Males) 1. Wampum, Cat. No. 61.2842 67 mm. 57 mm. 33 mm. .49

2. do. “ “ do 60 “ 51 “ 33 “ .55

3. do. “ “ do 50 “ 44 “ 25 " .50

(Females) 4. Wampum, Cat. No. 61.2842 47 “ 44 " 25* “ .53

5. Natrona, Cat. No. 61.4116 40 “ 36 “ 22 “ .55

6. Creekside, Cat. No. 61.3269 37 “ 31 “ 19 “ .51

The lowest diameter ever observed is in a specimen from West Fork River in
West Virginia, where it is forty-six percent of the length.

Soft parts identical with those of 0. subrotunda (See Ortmann, 1912, p. 328).
Glochidia figured by Ortmann, 1911, PL 89, fig. 15. They measure: 0.20 X 0.23
mm.

Breeding season: Glochidia have been found on Sept. 21, 1908, and again on
May 23, 1911; May 24, 1911; and May 27, 1908. On the latter date, discharge
has been observed. This indicates, that the form is hradytictic.

Remarks: The essential difference between this and the normal form is the
compression of the valves. It is true that the color is also often lighter (as pointed
out by Lea) but this does not hold good for the average of the Pennsylvanian
specimens. As is evident by the measurements given, the diameter of the shell
is less than in 0. subrotunda; but it must be noted, that at certain places the two
forms are found associated, and that they actually and insensibly pass into each
other. The dividing line at the diameter of sixty percent is entirely artificial,
and does not express natural conditions. It has been introduced, as in similar
cases, simply for the sake of convenience.

In the smaller creeks, var. levigata is found rather exclusively, while subrotunda
prevails in the Ohio. The transitional zone between them is precisely in the region
of Pittsburgh. Thus these forms must be regarded as varieties of one and the
same species.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Neville Island, Allegheny Co.

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Mahoning River, Mahoningtown, Lawrence Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Clarksville, Mercer Co.

Pymatuning Creek, Pymatuning Townsliip, Mercer Co.

Allegheny River, Natrona, Allegheny Co.; Godfrey, Armstrong Co.

Crooked Creek, Rosston, Armstrong Co.; Creekside, Indiana Co.

228

MEMOIRS OF THE CARNEGIE MUSEUM.

Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).^^

Other localities represented in the Carnegie Museum:

Lake-drainage:

St. Joseph River, Fort Wayne, Allen Co., Indiana (C. Goodrich).

Ohio-drainage:

Tuscarawas River; Ohio (Holland collection).

West Fork River, Lynch Mines, Harrison Co.; Lightburn and Weston, Lewis Co., West Virginia.

Little Kanawha River, Burnsville, Braxton Co., West Virginia.

North Fork Hughes River, Cornwallis, Ritchie Co., West Virginia.

Elk River, Sutton, Braxton Co., West Virginia.

Pocatalico River, Raymond City, Putnam Co., West Virginia.

Coal River, Sproul, Kanawha Co., West Virginia.

Mud River, Milton, Cabell Co., West Virginia.

Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.

Licking River, Farmer, Rowan Co., Kentucky.

Tennessee-drainage :

Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).

Elk River, Estill Springs, Franklin Co., Tennessee (H. H. Smith).

Hurricane Creek, Gurley, Madison Co., Alabama (G. H. Clapp, donor).

Flint River, Maysville, Madison Co., Alabama (H. H. Smith).

Paint Rock River, Paint Rock, Holly Tree, Trenton, and Princeton, Jackson Co., Alabama (H. H. Smith).
South Chickamauga Creek, Ringgold, Catoosa Co., Georgia.

Holston River, Holston Station, Grainger Co., Tennessee.

Distribution and Ecology (See fig. 23) : Type locality, Kentucky River (Rafin-
esque) .

As has been mentioned above, it is hard to trace the distribution of this form
in its relation to 0. subrotunda. Simpson (1900) gives only: “Ohio-drainage
and southern Michigan.”

According to my experience this is the small-stream-form of 0. subrotunda,
and wherever the latter tends to ascend into the tributaries of the larger rivers,
it turns into levigata. This is true in the Beaver-drainage, in the Allegheny and
Monongahela in Pennsylvania. But this form is not found in all the tributaries.
Upon the whole it is one of the rarer shells in our state. Rather compressed
shells are found down the Ohio for a certain distance, but they become scarcer and
scarcer. I have traced them down as far as Portland, Meigs Co., Ohio. But since
there are only isolated specimens among large numbers of typical subrotunda, I
have not taken the trouble of picking them out.

This form must have once existed in the Monongahela near Point Marion, Fayette Co., for
specimens found in an Indian garbage heap are this (Ortmann, 1909c, p. 13). It is abundant in the
upper Monongahela system (see below).

ORTMANN: monograph op the naiades of PENNSYLVANIA.

229

In the tributaries of the Ohio in West Virginia it appears to be the rule that
levigata is the form of the smaller streams. In Ohio aside from the Mahoning
River (Dean) this form is positively known to occur in the Tuscarawas River
(Dean, 1890, also Carnegie Museum), and it has been reported from the upper
Scioto, Columbus, Franklin Co. (Lea), and it has crossed over into the lake-drainage
(St. Joseph River).

Further, Call (1885) cites it from Tennessee in Elk River, Lincoln Co., but
says that it is identical with “circulus” That it is actually there, a little farther up,
is shown by specimens in the Carnegie Museum. It also is found in a number
of the tributaries of the Tennessee in northern Alabama and northern Georgia,
and has been detected in the Holston above Knoxville (very rare at this point).

Ecologically this form is in Pennsylvania and West Virginia distinctly sand-
loving. It is found in bars of sand or fine gravel, and is even not averse to shifting
sand.

Subgenus Pseudoon Simpson (1900).

Ortmann, 1912, p. 321; Simpson, 1914, p. 298.

Type Amhlema olivaria Rafinesque.

Only one species is found in Pennsylvania.

Obovaria (Pseudoon) olivaria (Rafinesque) (1820).

Obovaria ellipsis (Lea) Simpson, 1914, p. 299; Obovaria olivaria (Rafinesque)

Vanatta, 1915, p. 553.

Plate XIII, figs. 8, 9.

Records from Pennsylvania:

Stupakoff, 1894 (Allegheny Co.).

Ortmann, 1909&, p. 192.

Characters of the shell: Shell of medium size, thick and solid. Outline elliptical
or subovate, longer than high, strongly oblique. Anterior and lower margins
forming a regular curve, the lower margin curving up behind, and joining the pos-
terior margin in a curve, without forming an angle, but posterior end of shell more
narrowly rounded than the anterior. Upper margin convex, passing gradually
into the descending posterior margin. Beaks swollen, directed obliquely forwards,
incurved, and located much anteriorly, but generally a little behind the most
anterior part of the anterior margin, elevated somewhat above the hinge-line.
Beak-sculpture rudimentary, consisting of four or five fine bars, which are sinuate
in the middle. The posterior part of the bars is rudimentary and disappears upon

230

MEMOIKS OF THE CAENEGIE MUSEUM,

the posterior slope. There are specimens with well preserved beaks, where there
is hardly a trace of these bars. Valves rather convex, chiefly so towards the
beaks. No posterior ridge developed. No sculpture upon the disk.

Epidermis normally gueenish olive, lighter or darker, but shading to yellowish,
and in old specimens to brownish olive, but never dark brown. Color generally
rather uniform, and growth rests not marked by distinct concentric bands. In
some specimens, chiefly young ones, faint gveen rays are discernible, but rays are
mostly absent.

Hinge well-developed. Pseudocardinals heavy and solid, stumpy or slightly
elongated, one in right, two in left valve, not distinctly divergent, and becoming,
in older shells, subparallel to the laterals. Laterals moderately long, heavy.
Interdentum moderately developed. Beak-cavity not very deep. Dorsal muscle-
scars in the beak-cavity. Adductor-scars distinct and deeply impressed, chiefly
the anterior ones. Nacre silvery white.

Sexual differences present in the shell, but not very strongly marked. In
the male the lower margin curves up backward from about the middle, and the
posterior end of the shell is distinctly narrowed, sometimes almost bluntly pointed.
In the female the lower margin begins to ascend at a point back of the middle,
and curves up more broadly, so that the shell is less narrowed behind, and the
posterior end is broader and more evenly rounded. In consequence of this the
female shell appears higher and shorter. The males also grow to a size distinctly
larger than the females. However, there are females in which the characteristic
shape is poorly developed, and sometimes males are more broadly rounded behind,
so that it is not always possible to positively tell the sex by the shape of the shell.

L. H. D.

Size: 1. Industry, Cat. No. 61.3561 (gravid 9 ) 66 mm. 54 mm. 36 mm.

2. Cooks Ferry, Cat. No. 61.4420 (9 ?) 59 “ 47 “ 34 “

3. Industry, Cat. No. 61.3561 (9) 42 “ 32 “ 20 “

4. Toronto, Cat. No. 61.5444 (o') 76 “ 60 “ 40 “

5. Portland, Cat. No. 61.4776 (o') 70 “ 55 “ 36 “

6. do. “ “ do. (o') 60 “ 49 “ 31 “

Soft parts: Ortmann, 1912, p. 323. Glochidia, ibid., PL 19, fig. 11; Surber,
1912, PL 2, fig. 25. According to Surber they measure: 0.210 X 0.265; while
my figures are: 0.19 X 0.22 mm.

Breeding season: Gravid females were collected: Aug. 29, 1908 (eggs); Sept.
22, 1910 (glochidia); Sept. 24, 1910 (glochidia); and June 20 and 21, 1911 (glo-
chidia, discharging). Although these represent only a few dates, it is clearly shown.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

231

that this is a hradytictic form, breeding from August to June, Surber’s records also
confirm this (1912, p. 7) : February, May, June, and August, September, October,
November, December.

Remarks: A species easily recognized by the short-elliptical or ovate outline,
with the margins more regularly rounded (without angles) than in any other species,
by its oblique shape, with much anterior beaks, convex shell, and greenish-olive
color. The hinge-teeth are also quite peculiar.

In the outline and convexity of the shell there is much variability, the posterior
end being more broadly or more narrowly rounded, and the beaks being more or
less anterior. In old shells the beaks are generally more anterior, sometimes
even being at the anterior end. The color also is somewhat variable, and old
shells are often discolored and dull brown, but not dark brown. In young shells
the pseudocardinals are somewhat divergent and not so distinctly parallel to the
laterals.

Localities represented in the Carnegie Museum:

Ohio River, Industry and Cooks Ferry, Beaver Co., Pennsylvania.

Ohio River, Toronto, Jefferson Co., Ohio; St. Marj^s, Pleasants Co., West Virginia; Portland, Meigs Co.,
Ohio; Portsmouth, Scioto Co., Ohio.

West Fork White River, Riverside, Greene Co., Indiana (J. D. Haseman).

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).

Mississippi River, Moline, Rock Island Co., Illinois (P. E. Nordgren).

Kansas River, Lawrence, Douglas Co., Kansas (R. L. Moodie).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).

Distribution and Ecology (See fig. 23) : Type locality, Kentucky River (Rafin-
esque) .

One of the rarest species in Pennsylvania. It reaches the state only in the
Ohio River in Beaver and Allegheny Counties, and does not extend upstream
beyond Pittsburgh.

Farther down the Ohio, it is found only in this river, and does not go into the
tributaries in the state of Ohio (Sterki, 1907a), but it is rather common here. It
has a wider distribution in Indiana and Illinois. In Indiana (Call, 1896a and 1900)
it is practically all over the state, and crosses over into the lake-drainage (Maumee),
and goes even farther north into southern Michigan (Detroit, Grand, and Saginaw
Rivers, according to Walker, 1892, 1894, and 1898). It has been reported from
Lake Erie (Walker, 1913), and also from western New York, Niagara River and
Cayuga Lake (Marshall, 1895), and from the lower St. Lawrence in Canada (Ottawa,
Montreal, and Quebec, see Marshall, 1895; Bell, 1859; Whiteaves, 1863). How-
ever this northeastward expansion of the range is peculiar in its detail, and should

232

MEMOIRS OF THE CARNEGIE MUSEUM.

be studied more closely; the species has not been observed on the Pennsylvanian
shores of Lake Erie.

Westward, it is abundant in Illinois (Baker, 1906), chiefly in the larger rivers,
the Ohio, Wabash, Illinois, Spoon, and Kankakee, and also in the Mississippi,
where it is also found on the Iowa side, and goes up to southern Wisconsin (Lapham,
1860) and Minnesota (Grant, 1886, Holzinger, 1888).

Thence westward and southwest ward, records are at hand from Missouri
(Utterback, 1916), Kansas (Scammon, 1906). Simpson (1900) also mentions the
Arkansas River. The Carnegie Museum has it from Black River in northern
Arkansas.

South of the Ohio it is found in the Kentucky River (type locality) and in the
Cumberland (Wilson & Clark, 1914). It was reported from Nashville, Davidson
Co., Tennessee, by Marshall (1895). A single specimen is in the Carnegie Museum
from the Tennessee in northern Alabama. Farther up above Chattanooga it is
absent. I never saw it there, and it is absent in I^ewis’ list (1871).

As to its ecology I am able to say from personal experience, that it is a species
belonging to the shell-banks of the Ohio, in the deep channels, with strong, steady
currents. It is also in the smaller branches in (at low stages) shallow water, in
gravel. Scammon (1906) calls it ‘^a lover of water of moderate depth and of sandy
river-beds.”

Genus Actinonaias Crosse & Fischer (1893).

Nephronaias Crosse & Fischer; Ortmann, 1912, p. 324; Actinonaias Frierson, 1917,
p. 48.'"®

Type Unio sapotalensis Lea.

The only species known from Pennsylvania has been placed by Simpson in
Lampsilis. But whatever the final name of the genus may be, it is surely not
Lampsilis, because entirely lacking the characteristic structures found in the
female in that genus.

Actinonaias ligamentina (Lamarck) (1819).

Lampsilis ligamentina (Lamarck) and Lampsilis ligamentina nigrescens Simpson,

1914, pp. 79 and 82.

Plate XIV, figs. 5, 6.

Records from Pennsylvania :

Harn, 1891 (western Pennsylvania).

Stupakoff, 1894 (Allegheny Co.).

Simpson’s Nephronaias contains an assemblage of heterogenous species, belonging even to
different subfamilies. The nomenclature of this genus is still provisional, the structure of the type-
species being as yet incompletely known.

OETMANN: monograph of the naiades of PENNSYLVANIA.

233

Marshall, 1895 (Allegheny River, Warren Co.).

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.; Beaver River, Wampum,

Lawi’ence Co.).

Ortmann, 19096, p. 190.

Characters of the shell: Shell large, thick, and heavy. Outline subelliptical or
subovate, more or less elongated. Anterior margin rounded, lower margin and
upper margin together with the posterior margin forming rather regular curves,
joining posteriorly in a blunt point, situated below the horizontal middle line of
the shell. Beaks moderately swollen, little elevated, situated anterior to the
middle of the shell. Beak-sculpture poorly developed, consisting of a few fine,
indistinct bars, which have a tendency to be double-looped. Valves moderately
and rather uniformly convex; posterior ridge faint, indistinct, or obliterated. No
sculpture upon the disk.

Epidermis yellowish, greenish, or brownish-olive, often with more or less
distinct rays. Rays present chiefly in young individuals, broad and continuous,
sometimes sharply marked, sometimes obscure, covering the whole shell. In other
cases, there are hardly any traces of rays, and in old specimens the epidermis
is uniformly brown or blackish. Concentric bands of color are sometimes present.

Hinge well-developed. Pseudocardinals normally one in right, two in left
valve, but often additional ones are present. The normal pseudocardinals are
strong and heavy, ragged, subtriangular, divergent. Interdentum narrow, rather
long. Laterals long, strong, heavy. Beak-cavity moderate. Dorsal muscle-scars
in beak-cavity and upon the hinge-plate. Adductor-scars distinct, well impressed,
chiefly the one anterior. Nacre silvery white, but often discolored, in very rare
cases pinkish.^^^

Sexual differences present, but slight. In the male the lower margin is rather
evenly curved. In the female it is more curved out in its posterior part, so that
the poterior point of the shell is more elevated, and the whole posterior end of the
shell appears higher and more rounded. But in many cases, it is hard, or even
impossible, to tell the sex from the shape of the shell. Wherever the posterior
expansion is distinctly developed, we may be sure to have a female before us.

L. H. D.

Size: 1. Edinburg, Cat. No. 61.3491 ( 9 gravid) 155 mm. 95 mm. 61 mm.

2. Pulaski, Cat. No. 61.4059 ( 9 ) 147 “ 94 “ 63 “

3. Edinburg, Cat. No. 61.3491 (cf) 124 “ 76 “ 51 “

4. Cooks Ferry, Cat. No. 61.4791 (cf') 93 “ 65 “ 41 “

Such specimens have never been found in Pennsylvania, but I have a specimen with pinkish
nacre from Blennerhasset Island, near Parkersburg, West Virginia.

234

MEMOIRS OF THE CARNEGIE MUSEUM.

Soft parts (See Ortmann, 1912, p. 325). Glochidia: Lea (Obs. VI, 1858, PI. 5,
fig. 18) Ortmann (19116, PL 89, fig. 16). Surber, 1912, PI. 2, fig. 18. My measure-
ments are: 0.22 X 0.24; those of Surber: 0.220 X 0.260 mm.

Breeding season: Records for gravid females are rather complete, and cover
the period from Aug. 3 to Oct. 24, and from March 21 to May 20. This is a hrady-
tictic form with the interim in June and July. According to Surber (1912, p. 7),
glochidia are present from September to July, with the interim in August (in Iowa).

Remarks: Externally a form of simple appearance, without striking character-
istics. It may be recognized by its large size and heavy shell, of a rather regular
subelliptical outline, with more or less distinct, broad rays. Young shells might
be mistaken for Eurynia iris, or even for Lampsilis luteola, but these latter are
both more elongated, and differ somewhat in color.

A. ligamentina varies a good deal. The southern variety, gihba of Simpson,
is not represented in Pennsylvania. Humped specimens (the character of gihba),
are found, but they have the earmarks of cripples. The var. nigrescens (Simpson,
1914, p. 82) reported from Allegheny County, is merely an individual variation,
which is assumed occasionally by old shells, and does not deserve a name.

There are peculiar forms in Arkansas, which generally go by the name of liga-
mentina. They differ from the normal form in the color of the epidermis and of
the nacre. I have not made up my mind with regard to these, and in the following
list of localities they have been omitted, as has been the var. gibba. Only un-
doubted representatives of A. ligamentina are recorded.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio-drainage:

Ohio River, Smith’s Ferry, Sliippingport, Cook’s Ferry, and Industry, Beaver Co.; Beaver, Beaver Co.
(W. E. C. Todd); Dead Man’s Island, Shousetown, and Neville Island, Allegheny Co.; Coraopolis
(S. N. Rhoads) and Edgeworth (G. H. Clapp), Allegheny Co.

Little Beaver Creek, Cannelton, Beaver Co. (Miss Vera White).

’‘Beaver-drainage :

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Ell wood City, Lawrence Co. (G. H. Clapp & H. H. Smith).

Slipperyrock Creek, Wurtemberg, Lawrence Co.

Mahoning River, Mahoningtown, Coverts, and Edinburg, Lawrence Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.

Allegheny-drainage :

Allegheny River, Sandy Creek, Harmarville, and Natrona, Allegheny Co.; Braeburn, Westmoreland Co.;
Schenley, Aladdin, Godfrey, Johnetta, Kelly, Rosston, Mosgrove, Templeton, and Parkers Land-
ing, Armstrong Co.; Walnut Bend, Venango Co.; Tionesta and Hickory, Forest Co.; Warren,
Warren Co.

OETMANN: MONOGEAPH op the naiades of PENNSYLVANIA.

235

Conemaugh River, New Florence, Westmoreland Co.

Loyalhanna River, Idlepark, Westmoreland Co. (D. A. Atkinson).

Crooked Creek, Rosston, Armstrong Co.

French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co.
Connewango Creek, Russell, Warren Co.

Monongahela-drainage:

Monongahela River, Charleroi, Wasliington Co. (G. A. Ehrmann).

Youghiogeny River, Boston, Allegheny Co. (D. A. Atkinson).

Cheat River, Cheat Haven, Fayette Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Grand River, Cayuga, Haldimand Co., Ontario, Canada (C. Goodrich).

Sandusky River, Fremont, Sandusky Co., Ohio (C. Goodrich).

Maumee River, Defiance, Defiance Co., Ohio (C. Goodrich).

Raisin River, Monroe and Grape P. 0., Monroe Co., Michigan (C. Goodrich).

Huron River, Newport, Monroe Co., Michigan (C. Goodrich).

Grand Rapids, Kent Co., Michigan (Hartman collection).

Ohio-drainage:

Ohio River, Congo, Hancock Co., West Virginia; Toronto, Jefferson Co., Ohio; St. Marys, Pleasants
Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland, Meigs Co., Ohio; Ports-
mouth, Scioto Co., Ohio.

Tuscarawas River, Ohio (Holland collection).

West Fork White River, Riverside, Green Co., Indiana (J. D. Haseman).

Little Kanawha River, Grantsville, Calhoun Co., West Virginia (W. F. Graham).

Elk River, Shelton and Clay, Clay Co.; Gassaway, Braxton Co., West Virginia.

Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.

Licking River, Farmer, Rowan Co., Kentucky.

Mississippi and westward:

Kishwaukee River, Rockford, Winnebago Co., Illinois (P. E. Nordgren).

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection).

Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).i^*

Distribution and Ecology in Pennsylvania (See fig. 24) : A very common species
in western Pennsylvania, but restricted to the larger streams and some of the
larger tributaries. In the Ohio, Allegheny, and Monongahela, it is (or was) every-
where present, but in the two latter only up to a certain point. In the Allegheny
it goes up to Warren, and beyond, at least as far as Olean, Cattaraugus Co., New
York (Marshall, 1895), but is not found in the uppermost part in McKean Co.,
Pennsylvania. In the Monongahela it has been found as high up as the Cheat,

Mostly normal; but one specimen agrees with one of the Arkansas forms, with dark bold rays,
and pink nacre.

236

MEMOIRS OF THE CARNEGIE MUSEUM

but I never have seen it in the upper Monongahela (West Fork River) in West
Virginia. Likewise in the tributaries, it only goes up for a certain distance, and
then rather suddenly stops. This is the more remarkable, since it is, where present,
the prevailing species, outnumbering all other species combined, and then dis-
appears entirely sometimes within a few miles. In the Little Beaver I have never
seen it above Cannelton. It is plentiful at Edinburg in the Mahoning, but at
Hillsville, a few miles farther up, I saw only a single dead shell, and it is absent

• Actinonaias ligamentina.

in the Ohio part of this river (Dean, 1890). It is plentiful at Pulaski in the She-
nango, but above Sharon there is no trace of it. It goes into the Connoquenessing
to Ellwood City, and enters the mouth of Slipperyrock Creek, but is not found
farther up. It used to be found in the Kiskiminetas, but our material from this
stream is scanty. A dead shell was found in the Loyalhanna at Idlepark. In
Crooked Creek it occurs only at the mouth; in French Creek it extends up to
Cambridge Springs, and in Connewango Creek as far up as Russell.

Thus it is clear that A. ligamentina prefers the larger rivers. It is found here
in various environmental conditions, but is apparently best fitted for rough parts,
riffles with strong currents and heavy gravel and rocks. Baker (1898a) says that
it is found in muddy and sluggish rivers in soft mud, but this is decidedly not the
case in our region. CaU (1900) also calls it a “mud-loving species.” In the Ohio
below Pittsburgh and far down toward Cincinnati it is the form mainly composing
the shell-banks.

OETMANN: MONOGKAPH op the naiades of PENNSYLVANIA.

237

General distribution: Type locality, Ohio River (Lamarck).

That this species favors the larger rivers seems to hold good throughout the
rest of its range, as well as in Pennsylvania. It is found nevertheless in the north-
ern tributaries of the Ohio in Ohio, Indiana, and Illinois. In this region, it had a
chance to cross over into the lake-drainage, and is present in the Cuyahoga River
in Ohio (Dean, 1890, Dali & Simpson, 1895), in the Maumee-drainage in Ohio and
Indiana (Call, 1896a), and in the tributaries of Lake Michigan in Indiana (Call)
(See also our material). It is not found in the Michigan-drainage in Illinois,
although it ascends in the Desplaines River into the Chicago area (Baker, 1898a).
It occurs also in the southern part of Michigan (Walker, 1898) in the Lake Erie-
drainage, but is not in Lake Erie proper, and not in the lower St. Lawrence-drain-
age. From Illinois it goes northward into Wisconsin (occurring also in the
lake-drainage at Racine, Racine Co., according to Call (1885) and Milwaukee,
according to Lapham (1860) and Minnesota (Call, 1885; Grant, 1886; Holzinger,
1888) and here apparently crossing over into the northern drainage in Manitoba
(Simpson, 1900) reported from Roseau River by Dawson (1875). Call (1895)
mentions it from Dakota, but the particulars are not known. It certainly occurs
in Iowa, disregarding the Mississippi (See Call, 1895; Marshall, 1895; Geiser,
1910), and. the eastern part of Kansas (Scammon, 1906).

It is also found present south of this range and is known from the southern
tributaries of the Ohio in West Virginia and eastern Kentucky, from which speci-
mens are before me. It is commonly reported from the Cumberland and Tennessee-
drainages, but here the var. gibha turns up, and particulars as to the mutual rela-
tion of the distribution of the two varieties are lacking. All the material, without
exception, which I collected in the upper Tennessee region in Tennessee and Vir-
ginia represents the form gibba. Wilson & Clark (1914) say that in the Cumber-
land River “chiefly the southern mucket” {gibba) is found.

From Missouri southward into Arkansas ligamentina-iorms exist, and liga-
mentina has been reported from Missouri, for instance, by Utterback (1916) and
by Wheeler (1918) from the Ouachita in Arkansas. But, as has been stated,
different races (at least two of them) are found here, and these require further
study. In the Alabama-drainage, this type of Naiad seems to be entirely missing.

Simpson’s record from Ontario (1900) is vague, but the Carnegie Museum has it from Ontario
north of Lake Erie (Grand River).

238

MEMOIRS OF THE CARNEGIE MUSEUM.

Genus Amygdalonaias Crosse & Fischer (1893).

Ortmann, 1912, p. 327; Simpson, 1914, p. 306 (as subgenus of Plagiola).

Type Unio cognatus Lea.

The nomenclature of this genus must remain provisional, until we know the
anatomical structure of the type-species. Two species have been reported from
Pennsylvania.

Key to the Species of Amygdalonaias.

ai. Shell short and liigh, larger A. truncata.

a-i. Shell elongated and low, smaller A. donaciformis.

Amygdalonaias truncata (Rafinesque) (1820).

Plagiola elegans (Lea) Simpson, 1914, p. 307; Amygdalonaias truncata (Rafin-
esque) Utterback, 1916, p. 148.^^“

Plate XIV, fig. 7.

Records from Pennsylvania:

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co.).

Ortmann, 1909&, p. 192.

Characters of the shell: Shell rather small, moderately thick. Outline sub-
triangular-ovate, short and high. Anterior margin rounded, curving into the lower
margin, the latter nearly straight posteriorly, and often somewhat concave. Upper
margin short, curved, passing in a blunt angle or insensibly into the obliquely
descending posterior margin, which joins the lower margin in a more or less distinct
angle, so that the posterior end of the shell appears pointed. Beaks more or less
elevated, somewhat incurved, situated somewhat in front of the middle. Beak-
sculpture rudimentary, consisting of three or four fine bars, the first subconcentric,
those following double-looped; the posterior loop is triangularly pointed, and its
ascending part on the posterior slope is obsolete. Valves convex, more strongly
so in the anterior part of the shell, flattened upon the sides, and often with a faint
depression in front of the posterior ridge. The latter is distinct, often elevated
and keel-like towards the beaks. The posterior slope appears truncate, slightly
convex, flat, or even slightly concave.

Epidermis yellowish brown or greenish, mostly with well-developed rays.
Rays wide or narrow, straight. Very often there is a pattern of dark spots upon
the rays, which may be arrow-shaped, or wavy, and may extend over a large part

Vanatta (1915, p. 553) does not accept Rafinesque’s specific name on account of Unio truncata
Spengler (1793); however, this does not conflict, since Rafinesque called his species Truncilla truncata.
See also Walker, 1916, p. 45.

OKTMANN: MONOGEAPH of the naiades op PENNSYLVANIA.

239

of the surface. In older specimens the color-markings often become obscure, and
the epidermis is uniformly brown or blackish.

Hinge well-developed. Pseudocardinals two in left, one or two in right valve,
triangular, elevated and compressed, ragged. Interdentum quite narrow and
short. Laterals moderately long. Beak-cavity moderate. Dorsal muscle-scars
in beak-cavity. Anterior adductor-scars distinct and well impressed, posterior
ones faint. Nacre silvery white, rarely pinkish.

Sexual differences in the shell very indistinct. Specimens with a more distinct
furrow in front of the posterior ridge, and with the posterior section of the lower
margin concave, are generally males; while the opposite condition indicates the
female sex. In most cases it is very hard to determine the sex from the shell.

Size: 1. Neville Island, Cat. No. 61.1761,

2. Industry, Cat. No. 61.3569 (c?).

3. do. Cat. No. 61.3570 (cf).

L.

H.

D.

58 mm.

43 mm.

27 mm.

51 “

40 “

23 “

46 “

CO

22 “

No. 1 is one of the largest specimens at hand. No. 2 might be a female accord-
ing to the shape of the shell, but the soft parts had the male structure.

Soft parts (See Ortmann, 1912, p. 328). Glochidia: Lefevre & Curtis (1910,
p. 97, fig. 1; 1912, p. .146, fig. 3); Surber (1912, PL 2, fig. 30). They are very
small; 0.075 X 0.09 mm.

Breeding season: I found only one gravid female, with glochidia, on May
25, 1914. Lefevre & Curtis (1912, p. 141) list this species with the bradytictic
forms. Surber (1912, p. 6) states that gravid females with glochidia were found
in May and July.

Remarks: The species is easily recognized by its subtriangular shape, its sharp
posterior ridge, truncate posterior slope, and the peculiar color-pattern of the
epidermis. It only resembles the following species, but is distinguished from that
by the higher and shorter shell, which also gTows to a larger size. In addition this
species has a certain resemblance to the male of Truncilla triquetra both in general
shape as weU as in color. However Truncilla triquetra is a more elongated shell,
the posterior slope is broader, and more distinctly truncate, and the shell is more
inflated.

The posterior ridge of A. truncata is quite variable. In Pennsylvania it is
not so sharp as elsewhere. From Louisiana, for instance, I have specimens in which
it is keel-like. The form from Lake Erie is smaller and lighter, but very variable
in color. Having not been found on the Pennsylvanian shores of the lake, I shall
not here discuss this form.

240

MEMOIRS OF THE CARNEGIE MUSEUM.

Localities in Pennsylvania represented in the Carnegie Museum:

Oliio River, Shippingport and Industry, Beaver Co.; Neville Island, Allegheny Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Lake Erie, Vermilion, Erie Co., Ohio; La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).
Sandusky River, Fremont, Sandusky Co., Ohio (C. Goodrich).

Maumee and Erie Canal, Lucas Co., Ohio (C. Goodrich).

Ohio-drainage :

Ohio River, Toronto, Jefferson Co., Ohio; Parkersburg, Wood Co., West Virginia; Portland, Meigs
Co., Oliio.

West Fork White River, Riverside, Greene Co., Indiana (J. D. Ilaseman).

Wabash River, New Harmony, Posey Co., Indiana (A. A. Hinkley).

Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.

Tennessee-drainage :

Tennessee River, Florence, Lauderdale Co., Alabama (H. II. Smith).

Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).

Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).

Holston River, Hodges, Jefferson Co., Tennessee.

Clinch River, Solway, Knox Co.; Edgemoor and Clinton, Anderson Co.; Black Fox Ford, Union Co.;
Clinch River Station, Claiborne Co.; Oakman, Grainger Co., Tennessee.

West of Mississippi:

Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).

Marais des Cygnes River, Richhill, Bates Co., Missouri (W. 1. Utterback).

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

White River, Cotter, Baxter Co., Arkansas (A. A. Hinkley).

Black River (H. E. Wheeler) and Spring River (A. A. Hinkley), Black Rock, La^vrence Co., Arkansas.
Ouachita River, Arkadelpliia, Clark Co., Arkansas (H. E. Wheeler).

Verdigris River, Wagoner, Wagoner Co., Oklahoma (F. B. Isely).

Bayou Pierre, De Soto Par., Louisiana (L. S. Frierson).

Distribution and Ecology (See fig. 25) : Type locality, Ohio River (Rafinesque)
(Vanatta says “Falls of the Ohio”)-

In Pennsylvania this species is rare, and turns up only in the Ohio below
Pittsburgh. The few specimens I found there, were found in and below riffles,
probably washed out of shells-banks above them.

According to Simpson (1900) this form is generally found in the Mississippi-
drainage, but crosses over into the lake-drainage in southern Michigan, Lake
Michigan, and Lake Erie (Sterki, 1907a, Walker, 1913, and our localities). How-
ever it has not been found on the Pennsylvanian shores of Lake Erie.^®^ In Ohio

Call (1885) mentions it from western New York which extreme eastern extension might refer to
Lake Erie. Marshall (1895) quotes it from Buffalo, but from an entirely unreliable source.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

241

it is also occurs in tributaries to the lake, the Maumee, Tiffin, and Sandusky Rivers
(See Sterki). It is found in the Ohio-drainage in Ohio (Sterki), Indiana (Call,
1896a, 1900). It is widely distributed in Illinois (Baker, 1906). In the Missis-
sippi, it ascends from Illinois and Iowa to Wisconsin (Lapham, 1860) and as far as
Minnesota (Grant, 1886; Holzinger, 1888). It extends down the Mississippi and

Fig. 25.

■ Amygdalonaias truncata.

• Amygdalonaias donaciformis.

-F Plagiola lineolata.

to its western tributaries from Missouri (Utterback, 1916) to eastern Kansas
(Scammon, 1906), Arkansas (Call, 1895; Wheeler, 1918), Oklahoma (Carnegie
Museum), and western Louisiana (Frierson, 1899; Vaughan, 1893), and is found
in eastern Texas (Singley, 1893) and as far as Trinity River (Simpson). From the
southern tributaries of the Ohio records are scarce, but it is known from the Big
Sandy in Kentucky, from the Cumberland (Wilson & Clark, 1914), from Duck
River in Tennessee (Marshall, 1895), and from the Tennessee and its tributaries
in Alabama up to the Holston and Clinch in eastern Tennessee.

Lewis (1877) cites it from the Alabama-drainage, but this has never been
confirmed.

Amygdalonaias donaciformis (Lea) (1828).

Plagiola donaciformis (Lea) Simpson, 1914, p. 308.

Plate XIV, figs. 8, 9.

Records from Pennsylvania:

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co.).

Characters of the shell: Shell much like that of A. truncata, but distinguished
by the general shape, which is more elongated and less elevated. The posterior
ridge, although well marked toward the beaks, is not so sharp as in A. truncata,
and there is no depression in front of it. Color-markings of the epidermis of the

242

MEMOIRS OF THE CARNEGIE MUSEUM.

same general character, but the arrow-shaped spots are more prominent, and often
extend over large parts of the shell in a zig-zag pattern.

Male and female shells a little more distinct. In the male the posterior part
of the lower margin slopes up in a nearly straight line, rendering the posterior
end of the shell more sharply pointed, while in the female the lower margin is
more expanded and curved, making the posterior end more bluntly pointed. But
there are cases in which the difference is hardly noticeable. Sometimes in the
female the posterior part of the lower margin behind the expansion is slightly
concave.

L. H. D.

Size: 1. (No. locality) Cat. No. 61.787 (probably cf) 49 mm. 34 mm. 20 mm.

2. do. “ “ do. (probably 9) 41 “ 26 “ 18 “

Soft parts described, but rather unsatisfactorily, by Simpson in (Baker, 1898,
p. 91). I find that the soft jiarts agree completely with those of A. truncata. Glo-
chidia (See Surber, 1912, PI. 2, fig. 29; and Ortmann, 1914, p. 67). They also are
remarkable for their small size. Surber’s dimensions are: 0.060 X 0.063 mm.;
mine are: 0.05 X 0.06 mm.

Breeding season: Surber found eggs and glochidia in July. I received speci-
mens with glochidia collected on Aug. 18, 1912.

Remarks: This species is closely allied to A. truncata, and possibly may be
only a variety of it. But, as far as I can see, there are no intergrades between them.

This is the only species previously reported from western Pennsylvania, which
I have not collected myself. I have seen the specimens collected by Rhoads (now
in the Philadelphia Academy), and thus it is beyond doubt that it once existed in
our state. It may yet be found on the Pennsylvanian shores of Lake Erie.

Localities represented in the Carnegie Museum:

Lake-drainage:

Lake Erie, Vermilion (C. Goodrich) and Cedar Point (0. E. Jennings), Erie Co., Ohio; La Plaisance Bay,
Monroe Co., Michigan (C. Goodrich).

Miami and Erie Canal, Lucas Co., Ohio (C. Goodrich).

Ohio-drainage :

Ohio River, Parkersburg, Wood Co., West Virginia.

West Fork White River, Riverside, Greene Co., Indiana (J. D. Haseman).

Wabash River, New Harmony, Posey Co., Indiana (A. A. Hinkley).

Tennessee-drainage :

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).

Mississippi River and westward:

Mississippi River, Moline, Rock Island Co., Illinois (P. E. Nordgren).

ORTMANN; monograph of the naiades of PENNSYLVANIA. 243

Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).

Hinkston Creek, Columbia, Boone Co., Missouri (D. K. Greger).

Missouri River, St. Joseph, Buchanan Co., Missouri (W. I. Utterback).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).

Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).

Alabama-drainage:

Coosa River, Weduska Shoals, Shelby Co.; Riddles Bend, Cherokee Co., Alabama (H. H. Smith).

Distribution and Ecology (See fig. 25): Type locality, Ohio (Lea).

, In Pennsylvania this species has been found only in the Ohio in Allegheny
County. It seems to cover about the same range as A. truncata (See Simpson,
1900). It goes from Lake Erie down the Ohio, and its tributaries, and up the
Mississippi to Minnesota, and westward and southward down and across the Missis-
sippi to Kansas, Oklahoma, and eastern Texas. But in addition it has been re-
ported from Alabama (Simpson) and Coosa River (Call, 1885), and its presence in
the latter river is confirmed by specimens in the Carnegie Museum.

It has been referred to as a mud-loving shell by Baker (1898), and as an in-
habitant of sand-bars by Call (1900), and of the sandy and muddy beds of rivers,
avoiding smaller streams, by Scammon (1906).

Genus Plagiola Rafinesque (1820).^®^

Ortmann, 1912, p. 329; Simpson, 1914, p. 302.

Type Obliquaria lineolata Rafinesque.

Monotypic genus.

Plagiola lineolata (Rafinesque) (1820).

Plagiola securis (Lea) Simpson, 1914, p. 304; Plagiola lineolata (Rafinesque)
Vanatta, 1915, p. 553.^®^

Plate XIV, fig. 10, Plate XV, figs. 1, 2, 3.

Records from Pennsylvania:

Harn, 1891 (western Pennsylvania).

Clapp, 1895 (Allegheny Co.).

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co.).

Ortmann, 19095, p. 192.

Characters of the shell: Shell rather large, heavy and solid. Outline triangu-
larly-ovate, generally slightly longer than high. Anterior margin rounded, curving
into the lower margin, which is less convex, or nearly straight in its posterior part.

Not 1819. It was published in 1819 as “ nomen nudum.”

See also Walker, 1916, p. 45.

244

MEMOIRS OF THE CARNEGIE MUSEUM.

Upper margin short, convex, continued without any angle into the obliquely
descending posterior margin, which meets the lower margin in a distinct, but
rounded, lower posterior angle. Beaks moderately elevated above the hinge-line,
incurved, located in front of the middle of the shell. Beak-sculpture obscure,
consisting of two or three fine and faint, double-looped bars. Valves rather com-
pressed, moderately convex anteriorly, much more flattened on the sides, and
lieculiarly compressed towards the beaks. Posterior ridge very distinct, although
rounded, sharpest towards the beaks. Posterior slope, behind the ridge, narrow,
truncate, flat, almost concave towards the beaks, but slightly convex towards the
posterior end.

Epidermis greenish yellow to light brown, darker in old individuals, with more
or less distinct rays, which are rather narrow, dark brown or dark green, and are
generally broken into lunate or squarish, dark spots. In old specimens, the rays
may become indistinct, but traces of them and of the sjiots on them, are in most
cases preserved at least in jiart of the shell.

Hinge well-developed and heavy. Pseudocardinals two in left, one or two in
right valve, heavy, ragged. Interdentum well-developed, rather wide and flat.
Laterals heavy. Beak-cavity moderate. Dorsal muscle-scars partly in the beak-
cavity, partly upon the hinge-plate. Adductor-scars well-developed and rather
deep, chiefly the anterior ones. Nacre whitish.

Sexual differences of shell not very great, but generally well-marked. The
female shell remains considerably smaller than that of the male, is more inflated,
so that the posterior ridge is not so sharp, but there is hardly a difference in the
outline, except that the posterior end is more rounded.

L. H. D.

Size: (Males) 1. Cooks Ferry, Cat. No. 61.3565 118 mm. 91 mm. 51 mm.

2. do. “ “ do 107 “ 83 “ 49 “

3. do. “ “ 61.4421 94 “ 74 “ 38 “

(Females) 4. Cooks Ferry, Cat. No. 61.3565 (gravid) . 84 “ 63 “ 40 “

5. do. “ “ do. (gravid) . 80 “ 62 “ 38 “

6. Industry, Cat. No. 61.3568 (gravid) .... 70 “ 59 “ 37 “

The above measurements include the maximum records for either sex.

Soft parts (See Ortmann, 1912, p. 329). Glochidia (Lea, Obs. VI, 1858, PL 5,
fig. 8; Lefevre & Curtis, 1910, p. 97, fig. H, and 1912, p. 146, fig. H; Ortmann,
19116, PL 89, fig. 17; Surber, 1912, PL 2, fig. 14). My (maximum) measurements

Simpson (1900, p. 603) says that the female shell is swollen at the postbasal region. This is
hardly correct. The swelling of the female is noticeable chiefly over the lateral faces of the disk, but
very rarely in the postbasal region.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

245

are: 0,26 X 0.35 mm.; those of Lefevre & Curtis, 0.23 X 0.31; those of Surber:
0.230 X 0.330.

Breeding season: 1 have the following dates for gravid females: Sept. 10, 1908;
Sept. 12, 1908; Sept. 23, 1908; Sept. 24, 1910; Sept. 25, 1908; Oct. 3, 1908;
Nov. 4, 1914; Nov. 17, 1917. Furthermore: Febr. 6, 1911; June 14, 1911;
June 20, 1911; June 21, 1911; July 13, 1911, In every case glochidia were present,
and on the three days in June, discharging females were present. On July 13
only a single individual, having almost entirely discharged the glochidia, was found.
Thus the species is clearly hradytictic, and the breeding season probably begins in
August and lasts till June or July of the next year. The interim would include
part of July and August. Surber’s observations agree with this.

Remarks: This species is easily recognized and not liable to be confounded
with any other. The subtriangular, compressed shape, with a narrow posterior
truncation, and the peculiar color-pattern, are very distinctive characters. The
females are quite variable in shape and the compression of the shell is not so striking.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Smiths Ferry (W. F. Graham), Cooks Ferry, Shippingport, and Industry Beaver Co.; and
Neville Island, Alleghenj'- Co. (W. F. Graham).

Allegheny River, Godfrey and Kelly, Armstrong Co.

Monongahela River, Westmoreland Co., and Charleroi, Washington Co. (G. A. Ehrmann).

Other localities represented in the Carnegie Museum:

Ohio-drainage:

Ohio River, Toronto, Jefferson Co., Ohio; Beach Bottom, Brook Co., West Virginia (W. F. Graham);
Clarington, Monroe Co., Ohio; St. Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co.,
West Virginia; Portland, Meigs Co., Ohio; Portsmouth, Scioto Co., Ohio.

Cumberland- and Tennessee-drainages:

Cumberland River, Burnside, Pulaski Co., Kentucky (B. Walker, donor).

Tennessee River, Florence, Lauderdale Co., Alabama (H. II. Smith); Knoxville and Brabsons Ferry,
Knox Co., Tennessee.

Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).

Clinch River, Offutt, Anderson Co., Tennessee.

Mississippi river and westward:

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection).

Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).

Neosho River, Burlington, Coffey Co., Kansas (R. L. Moodie); Miami, Ottawa Co., Oklahoma (F. B.
Isely).

White River, Cotter and Norfolk, Baxter Co., Arkansas (A. A. Hinkley).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).

Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).

246

MEMOIRS OF THE CARNEGIE MUSEUM.

Alabama-drainage :

Coosa River, Wetumpka, Elmore Co.; Weduska Shoals and Wilsonville, Shelby Co.; Riverside and

Lock 4, St. Clair Co.; Minnesota Bend, Cherokee Co., Alabama (H. H. Smith).

Distribution and Ecology in Pennsylvania (See fig. 25) : In Pennsylvania this
species is found only in the Ohio, Allegheny, and Monongahela. In the Alle-
gheny it goes to southern Armstrong County, where it is quite rare. In the Monon-
gahela it is known only from Charleroi in Washington County, and on the opposite
side in Westmoreland County. Below Pittsburgh it is not a rare shell, and has
been found in considerable numbers in the riffles and the shell-banks in coarser or
finer gravel and in strong currents.

Farther down the Ohio this is a common shell in the shell-banks, and is regu-
larly taken by the clam-diggers.

General distribution: Type locality, Falls of the Ohio, at Louisville, Kentucky
(Rafinesque) .

This species primarily belongs to the large rivers of the interior basin. In
the Ohio its extreme upstream range reaches Pennsylvania. Farther dowm it is
mainly restricted to this river (Sterki, 1907a). In Indiana it also occurs in the
larger tributaries, the Wabash and West White Rivers (Call, 1896a, 1900) and has
crossed over, probably verj^ recently, into the Alaumee at Fort Wayne, Indiana
(Goodrich, 1914). This is the only known record from the lake-drainage. In
Illinois it is also found in the Wabash, Kaskaskia, and Illinois Rivers, in the latter
as high up as Peoria County (Forbes & Richardson, 1913), and to the Kankakee
(Baker, 1906). It ascends the Mississippi as far as Minnesota (Grant, 1886;
Holzinger, 1888). It is also found in the Cumberland River (Wilson & Clark,
19126 and 1914) and in the Tennessee it goes up according to Lewis (1871) to the
“Holston” near Knoxville, but it must be born in mind that the “Holston” of
Lewis is the Tennessee. I found it there myself and also in the lower Clinch in
Anderson County, but not in the Holston proper.

West of the Mississippi this species is found in Alissouri (Utterback, 1916)
in southeastern Kansas (Scammon, 1906), Arkansas and Oklahoma (See Call, 1895;
Wheeler, 1918; and the material in the Carnegie Museum) but its exact south-
western boundary has not been determined.

In addition it is present in the Alabama and Tombigbee drainages in Alabama
(Lewis, 1877; Call, 1885; Simpson, 1900; and material in Carnegie Museum)
and has reached Georgia in the Etowah River (Call, 1885).

Everywhere it seems to favor the “mussel-beds” in the larger rivers. Scam-
mon (1906) reports it from rocky riffles in Kansas, but says that it occurs in a

ORTMANN: monograph of the naiades of PENNSYLVANIA.

247

variety of locations, and Call (1900) says that it is found on sand, gravel, and
mud-bars, but preferably on the last.

Genus Paraptera Ortmann (1911).

Ortmann, 1912, p. 330.^®®

Type Unio fragilis Rafinesque.

Only the type-species is positively known to belong to this genus, although
it is probable that several others go with it. The type has been found in our state.

Paraptera fragilis (Rafinesque) (1820).

Lampsilis gracilis (Barnes) Simpson, 1914, p. 181; Lampsilis fragilis (Rafin-
esque) Vanatta, 1915, p. 552; Lasmonos fragilis (Rafinesque) Utterback,
1916, p. 152.

Plate XV, figs. 4, 5. 6.

Records from Pennsylvania:

Clapp, 1895 (Allegheny Co.)

Marshall, 1895 (Allegheny River, Warren Co.)^^®

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.)

Ortmann, 19095, pp. 192 and 202.

Characters of the shell: Shell large, bufthin. Outline subovate or subelliptical,
normally with a high posterior wing, which renders the outline subtriangular.
However, this wing may be obliterated, chiefly in old shells. There is also generally
a small anterior wing. Shell more or less symphynote at the wings. Anterior
margin rounded, united with the upper margin in a sharp angle. Lower margin
more or less regularly curved. Upper margin straight, elevated posteriorly,
forming the posterior wing and angle. Posterior margin obliquely descending,
sometimes somewhat concave just below the upper posterior angle, joining the
lower margin in a broad curve, so that there is no posterior angle. Beaks low,
hardly elevated above the hinge-line, located in the anterior portion of the shell.
Beak-sculpture faintly developed, rudimentary, consisting of three or four fine bars,
the first of which is subconcentric, the others double-looped, but only with the

Utterback (1916, p. 151) uses for this genus, the name of Lasmonos Rafinesque (1831). How-
ever, the type of Lasmonos (L. fragilis Rafinesque, 1831) is not Unio fragilis Rafinesque (1820), but
Unio leptodon Rafinesque (1820), for which Rafinesque proposed the subgenus Leptodea in 1820. Thus
Lasmonos is a synonym of Leptodea. If U. leptodon should prove to be congeneric with Paraptera fragilis,
then, of course, my Paraptera should give way, not to Lasmonos, but to Leptodea.

I doubt the correctness of this locality. In the Allegheny I found this species as far up as southern
Armstrong County, where it is very rare. No trace of it was seen above Oil City.

248

MEMOIRS OF THE CARNEGIE MUSEUM.

posterior loop distinct, while the anterior is obliterated. Valves rather compressed,
moderately and rather uniformly convex, slightly more flattened upon the sides.
No distinct posterior ridge. Posterior slope compressed, and even somewhat
excavated, chiefly when the upper posterior wing is well developed.

E})idermis smooth, light-colored, pale yellowish to pale greenish, sometimes
light olive-brown, with or without greenish rays. When present, the rays are
straight, continuous, greenish, ' rather narrow, and not well-defined. On the
posterior slope, the epidermis is lamellar, darker, often with indications of a few
stronger rays. Growth-rests generally distinctly indicated by dark concentric
bands.

Hinge poorly developed. Pseudocardinals two in left, one in right valve,
feeble, subtriangular and lamellar, compressed, often imperfect and represented
by mere ridges. Interdentum absent. Laterals long and thin, the lower one in
left valve often rudimentary. Beak-cavity shallow. Dorsal muscle-scars in an
oblique row in the beak cavity, but rather far from its extreme point. Adductor-
scars faintly impressed, the anterior ones more distinct than the posterior ones.
Nacre silvery white, in most cases with more or less pink, chiefly towards the
beak-cavity; sometimes entirely pinkish.

Sexual differences distinctly marked in the shell. In the male the lower
margin is regularly convex, curving up in its posterior half, and the posterior end
of the shell is rather narrowly rounded. In the female the lower margin is more
expanded in the postbasal region, so that a greater anterior section is descending,
while posteriorly it curves up in a very broad curve, rendering the posterior end of
the shell more broadly rounded. In the male the greatest height of the shell is
located more in the middle of the shell, in the female, it lies in its posterior portion.
The posterior expansion of the female is generally extremely thin.

L.

H.

D.

Size: (Males) I. Erie, Cat. No. 61.4807

. .116

mm.

78

mm.

38 mm.

2. Industry, Cat. No. 61.3549

. .100

C(

62

(i

33 “

3. Erie, Cat. No. 61.4808

, . . 99

64

((

32 “

4. Industry, Cat. No. 3549

, . . 67

42

a

21 “

(Females) 5. Erie, Cat. No. 61.4109

. . 100

(C

78

C(

32 “

6. do. Cat. No. 61.4807 (gravid)

.. 84

u

65

((

29 “

7. Edgeworth, Cat. No. 61.1360

. . 82

u

55

i(

23 “

8. Industry, Cat. No. 61.3549 (gravid) . .

. . 67

u

46

(C

21 “

The largest specimens at hand both from Lake Erie and the Ohio system are
given. This species does not reach its maximum size in Pennsylvania, and, as is
shown above, the specimens in Lake Erie are larger than those from the Ohio.

ORTMANN: monograph of the naiades op PENNSYLVANIA.

249

Soft parts (See Ortmann, 1912, p. 331). Glochidia (See Lefevre & Curtis,
1910, p. 97, fig. K, and 1912, p. 146, fig. K; Ortmann, 19115, PI. 89, fig. 19; Coker
& Surber, 1911, PI. 1, fig. 2; Surber, 1912, PI. 2, fig. 28). My measurements are:
0.08 X 0.09; those of Lefevre & Curtis: 0.07 X 0.09; those of Surber: 0.070 X
0.095 mm.

Breeding season: I have the following records for gravid females: Aug. 30,
1909 (beginning to have eggs); Sept. 8, 1908; Sept. 10, 1908; Sept. 11, 1913;
Sept. 17, 1913; Sept. 22, 1910; Sept. 25, 1908; Oct. 5, 1909. Then again in
spring: May 9, 1913; May 19, 1911; May 22, 1909; May 24, 1911; May 25,
1914; July 7, 1910; July 8, 1910; July 11, 1909. This is a hradytictic form, the
breeding season beginning in September. A specimen partly charged with eggs
was found as early as August 30. Toward the end of September glochidia are
found, but eggs have been observed as late as October 5. In spring, in May,
glochidia are present, and discharge has been observed on May 22 (in Lake Erie).
In the beginning of July the same has been found to be the case, and the latest
date for a discharging female, July 11, refers to a specimen from the Ohio River.
Thus part of July and August should be regarded as the “interim.” There does
not seem to be any difference between Lake Erie and the Ohio-drainage.

Surber found glochidia in October, November, January, June, July, and
August, and thus it appears that under certain conditions the seasons may overlap
in August.

Remarks: A species easily recognized by its subovate, compressed shape and
high posterior wing, and, if the latter is reduced, by its yellowish color, thin shell,
generally pink nacre, and weak or imperfect hinge-teeth. With regard to the
development of the wings there is great variability. Young specimens generally
have a well-developed, elevated, posterior wing, and a small anterior wing. But
in old specimens the wings may become lower, and are sometimes absent (in many
cases evidently broken off). This is chiefly true of specimens from the rivers
(Ohio), while the shells from Lake Erie preserve the wings better, even in large
individuals. This is undoubtedly due to the environment. The outline of the
shell varies greatly with the absence or presence of the wings being more regularly
elliptical, or subovate, or even sub triangular. There is also great variability in the
hinge, the pseudocardinals being sometimes distinct, sometimes very poorly
developed, knob-like, or like low ridges.

Contrary to what has been noted in other cases, the form from Lake Erie is
larger and finer than the Ohio form. This is undoubtedly due to the fact, that the
river-environment, as developed in western Pennsylvania, is not favorable to this

250

MEMOIRS OF THE CARNEGIE MUSEUM.

species, while the lake-environment suits it better. Otherwise there is no difference
between the two forms, except that the form from the Ohio river has on the average
a somewhat thicker shell and that the lake-form has a stronger tendency to pre-

• Paraptera fragilis.
+ Proptera alata.

serve the wings in old shells. But all these peculiarities due to environment are
more or less inconstant.

Localities in Pennsylvania, represented in the Carnegie Museum:

Ohio River, Smiths Ferry, Cooks Ferry, Industry, and Beaver, Beaver Co.; Dead Man’s Island (Q. T.

Shafer, A. T. Shafer, R. Foerster), and Edgeworth (G. H. Clapp), Allegheny Co.

Allegheny River, Aladdin, Godfrey, and Kelly, Armstrong Co.

Monongahela River, Westmoreland Co. (G. A. Ehrmann).

Lake Erie, Presque Isle Bay, Erie, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Lake Erie, Vermilion (C. Goodrich) and Cedar Point (C. Brookover), Erie Co., Ohio; La Plaisance Bay,
Monroe Co., Michigan (C. Goodrich).

Ohio-drainage:

Oliio River, Toronto, Jefferson Co., Ohio; Wheeling, Ohio Co., West Virginia (W. F. Graham); St.
Marys, Pleasants Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland, Meigs
Co., Ohio; Portsmouth, Scioto Co., Ohio.

West Fork White River, Riverside, Greene Co., Indiana (J. D. Haseman).

Little Kanawha River, Burnsville, Braxton Co., West Virginia.

Pocatalico River, Raymond City, Putnam Co., West Virginia.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

251

Tennessee-drainage :

Tennessee River, Florence, Lauderdale Co., Alabama (II. H. Smith).

Shoals Creek, Lauderdale Co., Alabama (H. H. Smith).

Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).

Tennessee River, Knox Co., Tennessee (Smith collection).

Nolichucky River, Chunns Shoals, Hamblen Co., Tennessee.

Holston River, Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill, Noeton, and Holston Station,
Grainger Co., Tennessee.

Clinch River, Edgemoor, Anderson Co.; Clinch River Station, Claiborne Co., Tennessee.

Powell River, Combs, Claiborne Co., Tennessee.

Mississippi River and westward:

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection); Moline, Rock Island Co.,
Illinois (P. E. Nordgren).

Hinkston Creek, Columbia, Boone Co., Missouri (D. K. Greger).

Kansas River, Lawrence, Douglas Co., Kansas (R. L. Moodie).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).

Spring River, Williford, Sharp Co., Arkansas (H. E. Wheeler).

Saline River, Benton, Saline Co., Arkansas (H. E. Wheeler).

Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).'

North Fork Canadian River, Weleetka, Okfuskee Co., Oklahoma (F. B. Isely).

Bayou Pierre, De Soto Parish, Louisiana (L. S. Frierson).

Alabama-drainage:

Buttahatchee River, Hamilton, Marion Co., Alabama (H. H. Smith).

Black Warrior River, Tuscaloosa, Tuscaloosa Co.; Squaw Shoals, Jefferson Co., Alabama (H. H. Smith).
Valley Creek, Toadvine, Jefferson Co., Alabama (H. H. Smith).

Coosa River, Wetumpka, Elmore Co.; near Yellow-Leaf Creek, Chilton Co.; Weduska Shoals and
Peckerwood Shoals, Shelby Co.; Coosa Valley and Riverside, St. Clair Co.; Minnesota Bend,
Cherokee Co., Alabama (H. H. Smith).

Distribution and Ecology in Pennsylvania (See fig. 26) : In Pennsylvania, this
species has two ranges. On the one hand it belongs to the three large rivers,
without, however, going into their tributaries. In the Allegheny it goes only as
far as southern Armstrong Co., where it is very rare; in the Monongahela, it has
been found only in Westmoreland Co. (opposite Charleroi). On the other hand
P. fragilis occurs in Lake Erie, where it is rather abundant in Presque Isle Bay.
Here it apparently finds most congenial conditions on the sandy shores in from
one to three feet of water; but it also has been obtained by the sand-sucker from a
depth of ten to fifteen feet.

In the Ohio and Allegheny I found this species chiefly in riffles. Here it is a
lively shell, crawling around frequently, and with a speed unusual in other shells.
The shells are here not very perfect, and often more or less injured or stunted.

General distribution: Type locality, Ohio River (Rafinesque) (Vanatta says:
“creeks in Kentucky”).

252

MEMOIES OF THE CAKNEGIE MUSEUM.

P. fragilis has an immense range, extending from central Texas (Singley, 1893)
north to the Red River of the North, and east and northeast over the whole Missis-
sippi and Ohio drainages, crossing over into the lake-drainage in Wisconsin, Ohio,
and Michigan, and going down the St. Lawrence to the Ottawa River (Call, 1885).
In this latter region it has extended also into Lake Champlain, and from western
New York (Marshall, 1895) through the Erie canal has reached the Hudson River,
thus entering the Atlantic watershed. In addition it is found in the Alabama and
Tombigbee drainages in Mississippi and Alabama (Lea, Obs. X, 1863; Conrad,
1836; Marshall, 1895). The latter part of the range is not given by Simpson.

It is not necessary to mention particular localities, but the western and south-
western boundaries, and also the northern boundaries, are rather indefinitely
known. The records from Kentucky and Tennessee were meagre hitherto, but
the species is present in the Cumberland (Wilson & Clark, 1914) and I found it
frequently in the upper Tennessee-drainage.

The ecological preferences of this species seem to be for large rivers and lakes,
and it avoids smaller rivers and creeks. Its absence from the Tuscarawas River
in Ohio, according to Sterki (1907a) and from the Beaver in Pennsylvania should
be noted. Where I found it in small streams, it was generally in eddies and pools.
But, as has been stated, it occurs also on riffles, but possibly it has been in such
cases washed out of the deeper and more quiet pools. Previous authors have
made similar observations (Call, 1895, 1900; Baker, 1898a; Scammon, 1906).

Genus Pkopteka Rafinesque (1819).^®^

Ortmann, 1912, p. 332; Simpson, 1914, p. 161 (as subgenus of Lampsilis).

lA^pe Unio alata Say.

Only one species in Pennsylvania.

Proptera alata (Say) (1817).^®^

Lampsilis alata (Say) Simpson, 1914, p. 162.

Plate XV, fig. 7; Plate XVI, figs. 1, 2.

Records from Pennsylvania:

Harn, 1891 (western Pennsylvania).

Stupakoff, 1894 (Allegheny Co.).

? Marshall, 1895 (Allegheny River, Warren Co.).^®^

This is the only one of the names proposed by Rafinesque in 1819, which has been properly
defined. The alate character of the shell has been mentioned, and a type (alata) has been given.

158 Not 1816.

155 1 have never seen a trace of this shell above Oil City, although it is not easily overlooked, and

ORTMANN: monograph of the naiades op PENNSYLVANIA. 253

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.).

Ortmann, 19096, pp. 191 and 202.

Characters of the shell: Shell very large, moderately thick. Outline subovate,
but generally with a high posterior wing, which makes the outline subtriangular.
This wing is very variable, but rarely (in old shells) indistinct. Sometimes there
is an indication of a small anterior wing. Shell symphynote at the wings. Anterior
margin rounded. Lower margin slightly convex, or almost straight posteriorly.
Upper margin straight, ascending posteriorly, and forming with the posterior
margin, the elevated wing-like angle. Posterior margin obliquely descending,
slightly concave just below the angle, or straight, and passing in a broad curve
into the lower margin, so that there is no lower posterior angle to the shell. Beaks
low, hardly elevated above the hinge-line, located in the anterior section of the
shell. Beak-sculpture indistinct, feeble, consisting of three or four fine bars, the
first subconcentric, the following ones double-looped, with the anterior loop almost
effaced. Valves rather compressed, gently convex, flattened upon the sides; no
posterior ridge, but shell in this region broadly rounded, with one or two fine
radiating, elevated lines. Posterior slope compressed or slightly excavated, ele-
vated into the posterior wing.

Epidermis thick, dark, in young specimens sometimes dark greenish, but
generally dark brown to black, without any rays. Mere traces of rays are rarely
visible in young specimens. Growth-rests are generally not marked by darker color.

Hinge well-developed. Pseudocardinals two in left, one or two in right valve,
moderately thick, triangular, ragged. No interdentum. Laterals long, generally
curved, moderately strong. Beak-cavity shallow. Dorsal muscle-scars in an
irregular, almost vertical row in the beak-cavity. Adductor-scars distinct, the
anterior impressed, the posterior less so.

Nacre always purple, generally very dark purple, rarely light purple, and in
one case I have seen a pale salmon-color, whitish toward the margins of the shell.

Sexual differences present in the shell, but less marked than in Paraptera
fragilis. The female is slightly more swollen in the posterior part of the shell,
and the lower margin is more broadly rounded in the postbasal region. These
differences, however, are slight, and not always reliable. Further, it seems that
the female does not attain the extreme size of the male.

if no living shells are found, dead ones are very conspicuous. Its uppermost limit in the Allegheny is
Templeton in Armstrong Co., and above Pittsburgh it is in the Allegheny altogether a rare shell.

254

MEMOIRS OF THE CARNEGIE MUSEUM.

L. H. D.

Size: (Ohio-form) 1. Wiley, Cat. No. 61.3537 (sex ?) 165 mm. 121 mm. 57 mm.

2. Godfrey, Cat. No. 61.4107 (cf) 157 “ 112 “ 44 “

3. Industry, Cat. No. 61.4416 (cf) 118 “ 84 “ 33 “

4. Industry, Cat. No. 61.3538 (9 gravid). . . 118 “ 79 “ 38 “ ,

5. Cooks Ferry, Cat. No. 61.3540 ( 9 gravid) . 101 “ 78 “ 35 “

(Lake Erie-form) 6. Erie, Cat. No. 61.4102 (9 gravid) 91 “ 68 “ 36 “

7. do. Cat. No. 61.3267 (cf) 91 “ 65 “ 32 “

8. do. Cat. No. 61.4816 (9 ) 87 “ 69 “ 31 “

9. do. Cat. No. 61.4817 (cT) 75 “ 58 “ 25 “

Soft parts (See Ortmann, 1912, p. 333). Glochidia: Lea (Obs. VI, 1858, PL 5,
fig. 25); Lefevre & Curtis, 1910, p. 97, fig. D, and PL 4, fig. 25; Ortmann, 19116,
PL 89, fig. 18; Coker & Siirber, 1911, PL 1, fig. 3; Lefevre & Curtis, 1912, p. 146,
fig. D; Surber, 1912, PL 1, fig. 8. My measurements of the glochidia are: 0.20 X
0.38; Lefevre & Curtis give: 0.23 X 0.41; and Surber: 0.220 X 0.380 mm.

Breeding season: The following dates for gravid females are at hand: June
24, 1909 (eggs); August 29, 1908; Aug. 30, 1909; Sept. 6, 1914; Sept. 8, 1908;
Sept. 10, 1908; Sept. 15, 1913; Sept. 17, 1908; Sept. 17, 1913; Sept. 23, 1908;
Sept. 25, 1908; Sept. 28, 1911; Oct. 3, 1908. Then again in spring: May 21,
1909; May 22, 1914; May 23, 1914; May 24, 1911 (discharging); July 7, 1910
(discharging); July 8, 1910 (discharging); July 11, 1909 (discharging).

The species seems to breed all the year round, and the breeding seasons appear
to overlap in June and July. Eggs have been found as early as June 24 (at Kelly)
but this might be an exceptional case. At the end of August fully developed
glochidia are present. On the other hand discharging females have been found
as late as July 11. These extreme dates refer to the Ohio-drainage. In Lake
Erie the time for discharging is the same, and I found females in this condition in
May as well as in the beginning of July. The beginning of the breeding season
has not been observed in the lake.

Remarks: This is a species easily recoguized by the large, winged shell, with
blackish epidermis, and purple nacre. In its external shape it much resembles
Lasmigona complanata (Barnes), while on the inside of the shell (hinge and nacre)
it is very different from it.

Proptera alata varies greatly in outline, chiefly on account of the variability of
the wing. In old specimens, sometimes also in young ones, the wing is almost
absent, and often it appears as if broken off in life.

The form from Lake Erie is distinguishable from the Ohio-form. It is smaller,
darker brown (not black), has a lighter nacre, and is on the average slightly more
swollen. A greater regularity of the growth rests (as is usual in lake-shells) is

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255

not evident in this case. If it should be considered desirable to distinguish the
Lake Erie form taxonomically, we should bear in mind that the type-locality for
U. alatus is Lake Erie, and that the lake-form should retain this name. The form
from the Ohio-drainage should then be known as P. alata megaptera (Rafinesque)
(1820).

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Shippingport, Cook’s Ferry, and Industry, Beaver Co.; Dead Man’s Island (Q. T. Shafer &
A. T. Shafer), Coraopolis (S. N. Rhoads), Neville Island, and Edgeworth (G. H. Clapp), Alle-
gheny Co.

Allegheny River, Natrona, Allegheny Co.; Aladdin, Godfrey, Johnetta, Kelly, and Templeton, Arm-
strong Co.

Little Beaver Creek, Cannelton, Beaver Co. (Miss Vera White).^®“

Monongahela River, Westmoreland Co., and Charleroi, Washington Co. (G. A. Ehrmann).

Dunkard Creek, Wiley, Greene Co.

Lake Erie, Presque Isle Bay, Erie, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Lake Erie, Vermilion (C. Goodrich), Cedar Point (C. Brookover), and Sandusky Bay, Cedar Point (0. E.

Jennings), Erie Co., Ohio; La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).

Maumee River, Roche de Boeuf Rapids, Lucas Co.; Defiance, Defiance Co., Ohio (C. Goodrich).

Ohio-drainage :

Ohio River, Congo, Hancock Co., West Virginia; Toronto, Jefferson Co., Ohio; Wheeling, Ohio Co.,
West Virginia (W. F. Graham); 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. (W. F. Graham); Burnsville, Braxton Co., West Virginia.
Pocatalico River, Raymond City, Putnam Co., West Virginia.^®i
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).

Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).

Tennessee River, Brabsons Ferry, 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, Noeton, and
Holston Station, Grainger Co., Tennessee.^®^

I have seen a dead shell in Little Beaver Creek at Smith’s Ferry, Beaver Co.

I have seen dead specimens in Elk River at Shelton and Clay, Clay Co., and at Sutton, Braxton
Co., West Virginia.

I have seen a dead specimen in the North Fork of the Holston at Rotherwood, Hawkins Co.,
Tennessee.

256

MEMOIRS OF THE CARNEGIE MUSEUM.

Clinch River, Solway, Knox Co.; Edgemoor and Offutt, Anderson Co.; Clinch River Station, Claiborne

Co.; Oakman, Grainger Co., Tennessee; Clinchport, Scott Co., Virginia.

Powell River, Combs, Claiborne Co., Tennessee.

Mississippi and westward:

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection); Moline, Rock Island Co.,

Illinois (P. E. Nordgren).

Osage River, Warsaw, Benton Co., Missouri (W. I. Utterback).

Bull Creek, Miami Co., Kansas (C. Goodrich, donor) (Osage-drainage).

Kansas and Wakarusa Rivers, Lawrence, Douglas Co., Kansas (R. L. Moodie).

Distribution and Ecology in Pennsylvania (See fig. 26) : This species has two
ranges in Pennsylvania: one in the Ohio system; the other in Lake Erie. In the
latter it is found in Presque Isle Bay, but is comparatively rare.

In the Ohio system, it belongs to the three large rivers, the Ohio, Allegheny
and Monongahela, and ascends the Allegheny to Armstrong Co., but is rare there:
Marshalhs record from Warren County should be disregarded. In the Monon-
gahela, it is known from the neighborhood of Charleroi, but must once have gone
farther iq), to near the West Virginia state-line, for it is found (although very
rarely) in the lower jiart of Dunkard Creek in Greene Co. There is only one
other instance known where it enters a small tributary, this is Little Beaver Creek
in Beaver County, and there also it is very rare. None of the other tributaries of
the Ohio system contains this species, which is most remarkable especially in the
case of the Beaver River, where this species has never been found.

In the Ohio below Pittsburgh P. alata is common, sometimes even very abun-
dant, and here it is found in riffles, in fine and often rather coarse gravel. It is a
rather active species, crawling around a good deal. It is also found on the shell-
banks at Industry and Shippingport, associated with the other bank-forming
species, in gravel and strong, steady currents.

In Lake Erie (Presque Isle Bay), it is found in the characteristic sand of the
north shore of the bay, in from two to three feet of water, and seems to prefer the
open shores (not covered and fringed by rushes).

General distribution: Type locality, Lake Erie (Say).

The centre of distribution is in the northern section of the interior basin, in
the Mississippi and Ohio Rivers, and their tributaries. In the Ohio its upper
boundary has been located in western Pennsylvania. Like Paraptera fragilis,
it passes northward into the northern drainage systems of the Red River of the
North, of Hudson Bay and of the St. Lawrence. In the latter it is found prin-
cipally in the lower lakes and their tributaries, as far as Ottawa (Bell, 1859; Whit-
eaves, 1863) and it also goes into Lake Champlain, Lake George, and the Hudosn-

ORTMANN: monograph of the naiades of PENNSYLVANIA.

257

drainage in New York, Waterford, Saratoga Co. (De Kay, 1843) Troy, Rensselaer
Co. (Aldrich, 1869). Compare also Marshall (1895).

Westward it extends to eastern Kansas (Scammon, 1906), and is also present
in the southern tributaries of the Ohio in West Virginia, Kentucky, Tennessee
(Marshall, 1895; Lewis, 1871; Wilson & Clark, 1914; Carnegie Museum) and
northern Alabama (Call, 1885; Carnegie Museum). In the Clinch River it
reaches Virginia.

Within the range thus indicated, this species seems to be present chiefly in
the larger rivers, but sometimes it is found in rather small streams.^®" Judging
from the Ohio below Pennsylvania, it is a species of the shell-banks in the deep
channels, and is taken in great numbers by the clam-diggers, but rejected as useless.
According to Baker (1898a) Call (1900) and Scammon (1906) it lives in the larger
lakes and rivers on muddy bottoms, but it certainly is not often found in mud in
Pennsylvania, where it occurs mostly in gravel.

Toward the south P. alata just reaches Kansas and (in the Tennessee) northern
Alabama. Beyond these stations, it is represented by allied, but distinct species
or forms. The true P. alata is positively missing from the Alabama-drainage.
Simpson (1914, p. 164) reports a form from this region, which is also represented
in the Carnegie Museum from the Coosa River, called var. potilsoni (Conrad).
This indeed closely resembles P. alata, but in my opinion is a variety of P. purpurea
(Lamarck). I cannot go into details here, but we have in this case probably to
deal with an interesting instance of ‘Yonvergency.”

Genus Toxolasma Rafinesque (1831).

Carunculina Ortmann, 1912, p. 337 (subgenus of Eurynia)] Carunculina Simpson,

1914, p. 148 (subgenus of Larnpsilis); Toxolasma, Frierson, 1914, p. 7;

Carunculina (genus) Ortmann, 1914, p. 68.

Type Unio lividus Rafinesque.

Only one species is known from Pennsylvania.

Its absence in the Tuscarawas River in Ohio has been noticed by Sterki (1907). This corre-
sponds to its absence in the Beaver River in Pennsylvania.

This is the Tennessee-Cumberland form of U. glans Lea. Particulars have been published else-
where.

258

MEMOIRS OF THE CARNEGIE MUSEUM.

Toxolasma parvum (Barnes) (1823).

Lampsilis parva (Barnes) Simpson, 1914, p. 151.

Plate XVI, fig. 3.

Records from Pennsylvania :

Harn, 1891 (western Pennsylvania).^®^

? Rhoads, 1899 (Beaver River, Wampum, LawTence Co.)^®®

Ortmann, 19096, p. 191.

Characters of the shell: Shell small, but rather solid. Outline subelliptical,
with the anterior and posterior ends almost uniformly and broadly rounded.
Lower margin gently convex or nearly straight, and almost parallel to the upper
margin. Beaks more or less inflated, slightly elevated above the hinge-line, located
anterior to the middle of the shell. Beak-sculpture distinct, consisting of five or
six comparatively strong, subconcentric bars, of which the later ones are sub-
angular behind. Valves more or less convex, often considerably inflated (chiefly
in the female), flattened upon the sides. No distinct posterior ridge. Posterior
slope flattened and somewhat compressed.

Epidermis thick, dark, greenish, brown, or blackish, without rays. Growth-
rests not indicated by concentric bands.

Hinge well-developed. Two pseudocardinals in left, one in right valve, tri-
angularly projecting, with rough edges. Interdentum absent. Laterals straight,
rather long, moderately strong. Beak-cavity moderately deep. Dorsal muscle-
scars in beak-cavity. Anterior adductor-scars distinct and well impressed, pos-
terior ones less so. Nacre silvery white.

Sexual differences present in the shell, but not very pronounced. As a rule
the male shells are less swollen, while the females are much more so (almost sub-
cylindrical). In the male the posterior end of the shell is narrower, sometimes
almost pointed, while in the female it is more broadly rounded and very blunt;
but there is great variability in this respect, and only in the older shells are these
differences unmistakable.

L. H. D.

Size: 1.

Conneautlake,

Cat. No. 61.3534

32

mm.

19 mm.

15 mm.

2.

do.

Cat. No. 61.4101

30

a

18 “

14 “

3.

do.

“ “ do

27

u

17 “

11 “

4.

do.

“ “ do

23

((

13 “

9 “

Nos. 2, 3, and 4 are gravid females, and No. 1 also has the shape of a female.

1®® Unfortunately Harn does not give exact locality, so that this record cannot be controlled.
i®6 Very likely incorrect, founded upon confusion with Eurynia fabalis.

1®’ I have a few southern specimens, wliich have a faint pinldsh blush.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

259

Soft parts (See Ortmann, 1912, p. 338). They have been figured by Lea, Obs.
VII, 1860, PL 29, fig. 102. As to the possible hermaphroditism of this species, see
Utterback (1916, p. 165). This question is yet rather obscure. Glochidia:
Lea (Obs. XIII, 1874, PL 21, fig. 3); Surber (1915, fig. 3); Utterback (1916, p.
165). I have seen the glochidia of specimens from Arkansas. They are subovate,
higher than long, 0.18 X 0.20 mm. Surber gives: 0.17 X 0.20. In Utterback’s
figures (0.175 X 0.100) apparently is a misprint.

Breeding season: In Pennsylvania I found three gravid females with eggs
on June 17, 1909, which would indicate a very early beginning of the season. But
on the other hand I have females from Arkansas with fully developed glochidia
collected June 26, 1911, and May 19, 1911. A specimen from Indiana, collected
Aug. 9, 1912, had eggs. Surber’s glochidia came from a specimen collected July
20, 1911. This is rather confusing, and additional data are much to be desired.

Remarks: The chief characteristics of this species are the small size, rather
regular subelliptic outline, the inflated valves, dark epidermis, and beak-sculpture.
In Pennsylvania it apparently has been confounded with Eurynia f abatis, which
resembles it in its small size, but is more compressed, is more pointed behind, has
much heavier hinge teeth, and generally more or less distinct rays.

Localities represented in the Carnegie Museum:

Conneaut Outlet, Conneautlake, Crawford Co., Pennsylvania.

Lake Erie, Put-in-Bay, Ottawa Co., Ohio (C. Goodrich).

Crane Creek, and Ten Mile Creek, Ottawa Park, Lucas Co., Ohio (C. Goodrich).

Grand Reservoir, Mercer Co., Ohio (C. Goodrich).^®*

Tuscarawas River, Ohio (Holland collection).

Scioto River and Ohio Canal, Columbus, Franklin Co., Ohio (Smith collection).

Wabash River, Mercer Co., Ohio (C. Goodrich); New Harmony, Posey Co., Indiana (A. A. Hinkley).
Big Creek, Solitude, Posey Co., Indiana (A. A. Hinkley).

Hihkston Creek, Columbia, Boone Co., Missouri (D. K. Greger).

Saline River, Benton, Saline Co., Arkansas (H. E. Wheeler).

Ouachita River, Arkadelphia, Clark Co., and mouth of Cove Creek, Hot Springs Co., Arkansas (H. E.
Wheeler).

Big Deceiper Creek, Gum Springs, Clark Co., Arkansas (H. E. Wheeler).

Malvern Creek, Malvern, Hot Springs Co., Arkansas (H. E. Wheeler).

Slough, Tulsa, Tulsa Co., Oklahoma (F. B. Isely).

Distribution and Ecology (See fig. 27): Type locality, Fox River, Wisconsin
(Barnes) .

In Pennsylvania this species is known from but one locality, the outlet of
Conneaut Lake. Here I found it in small numbers at a point where a small run

168 This drains originally into the Maumee, but now into the Wabash at high water.

260

MEMOIRS OF THE CARNEGIE MUSEUM.

emiities into the outlet, the water of the former being backed up and transformed
into a quiet pool. It lives here in mud.

This species has been reported from western New York in the Erie Canal in
Onondaga Co., and in the Genesee Canal (Marshall, 1895). It is possible that
these localities may later be discovered to be connected with the Pennsylvanian

• Toxolasma parvum.
m Eurynia fabalis.

+ Eurynia iris.

X Eurynia iris novi-eboraci.

locality, and this in turn with the western localities, when the species is looked for
at favorable habitats (canals, sloughs, etc.). In the state of Ohio it is not rare,
and is found in both drainages, and also in Lake Erie (Sterki, 1907a; Walker,
1913, p. 21). It is known in the lake-drainage in southern Michigan (Walker,
1898), and is found practically all over Indiana (Call, 1896a) and Illinois (Baker,
1906). Here it does not enter the lake-drainage, except in the Maumee River
(Call). It is known from Fox River, Wisconsin, from southern Minnesota (Call,
1885; Grant, 1886; Holzinger, 1888), from Iowa (Pratt, 1876; Witter, 1878;
Call, 1895; Marshall, 1895; Geiser, 1910). Thence it extends southward, and is
found in Missouri (Utterback, 1916), in Kansas (Scammon, 1906), Arkansas (Call,
1895; Wheeler, 1918; see also Carnegie Museum), Oklahoma, and as far south
as Texas (Singley, 1893). However, in this region a closely allied species, T.
texasense (Lea) turns up, and this seems to gradually replace it (see Simpson,
1900, p. 564, footnote 2) and possibly the two forms intergrade. According to
Vaughan (1893) they run into each other in northwestern Louisiana.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

261

South of the Ohio in Kentucky and Tennessee definite records are scarce.
Wilson & Clark (1914) give it from Stones River in Tennessee, a tributary of the
Cumberland. Call reports it (1885) from the Tennessee-drainage near Tuscumbia,
Alabama, and from Jackson, Hinds Co., Mississippi. But in this region and also
in Alabama it is supposed to be replaced by several other species, the standing of
which is very doubtful.

According to Baker (1898a) and Scammon (1906), this is a species living
buried in mud, in sluggish streams, canals, etc. This is supported by my observa-
tions in Pennsylvania.

Genus Eurynia Rafinesque (1820).

Ortmann, 1912, p. 336.^®^

Type Unio recta Lamarck.

Four species, and one variety are found in Pennsylvania. They fall into two
subgenera.

Key to the Subgenera of Eurynia.

ai. Inner edge of mantle in front of the branchial opening in the female with irregular, larger or smaller
papillae, generally rather distant from each other. Shell small or of medium size, subovate or
subelliptical, not very long (hardly over twice as long as high). Glochidia subspatulate, con-
siderably higher than long Subgenus Micromya.

02. Inner edge of mantle in front of branchial opening in the female with a long row of quite regular,
uniform, smaller or larger papillae, which stand rather crowded. Shell of medium or large size,
subelliptical, elongated, distinctly over twice as long as high. Glochidia subovate, only slightly
higher than long Subgenus Eurynia.

Subgenus Micromya Agassiz (1852).

Ortmann, 1912, p. 337.^^°

Type Unio f abatis Lea.

Two species and a variety in Pennsylvania.

Key to the Forms of Micromya.

tti. Shell very small. Hinge-teeth comparatively thick and heavy. Rays faint, broad or fine, often

somewhat undulating E. (M.) jahalis.

a 2. Shell larger, up to middle size. Hinge-teeth thin and delicate. Rays more or less distinct, not
undulating, but often interrupted.

fei. Rays straight and continuous, often indistinct E. (M.) iris.

1)2. Rays interrupted, forming blotches, generally well-developed E. {M.) iris novi-eboraci.

Eurynia (as subgenus of Lampsilis) Simpson, 1914, p. 60, is only in part identical with Eurynia
in our sense.

The genus Micromya of Simpson (1914, p. 32) contains two species, of which only the first
(fabalis) belongs here. The other species of Micromya in our sense stand under Lampsilis Simpson.

262

MEMOIKS OF THE CAENEGIE MUSEUM.

Eurynia (Micromya) fabalis (Lea) (1831).

Micromya Jabalis (Lea) Simpson, 1914, p. 33.

Plate XVI, figs. 4, 5.

Records from Pennsylvania:

Ham, 1891 (western Pennsylvania).

Marshall, 1895 (Allegheny River, Warren Co.).

Ortmann, 19096, p. 188.*^^

Characters of the shell: Shell small, but comparatively thick. Outline sub-
elliptical to subovate, rather short, generally a little less than twice as long as high.
Anterior and posterior ends rounded. Upper and lower margins more or less
curved, the lower sometimes nearly straight, or even concave in the middle. Beaks
not much elevated above the hinge-line. Beak-sculpture fine and rudimentary,
consisting of four or five bars, which are double-looped and interrupted in the
middle. The posterior loop is represented by a mere tubercle. Valves more or
less convex, flattened upon the sides. No distinct posterior ridge. Posterior
slope slightly convex.

Epidermis yellowish, or light or dark green. Generally covered all over with
green rays, but the rays may be absent on the anterior section. The rays are
not very sharp, and are broader or narrower, but on the posterior section of the
shell they are generally rather narrow, and very often more or less undulating.
On the posterior slope they are indistinct. Old shells often incline to a uniform
dark green, brown, or blackish color, with very indistinct rays. Growth-rests
obscure.

Hinge well-developed. Pseudocardinals two in left, one or two in right valves,
comparatively heavy, thick, stumpy, and somewhat crenulated. Interdentum
absent or narrow. Laterals moderately long, thick and heavy, especially the one
in the right valve, which stands upon a heavy basal plate. Beak-cavity shallow.
Dorsal muscle-scars in the beak-cavity. Adductor-scars distinct, and rather
deeply impressed, especially the one anterior. Nacre silvery white, sometimes
faintly salmon or pinkish toward the beak-cavity ; more or less iridiscent posteriorly.

Sexual differences rather well marked. The male shell is not much swollen,
and has a long-ovate outline, with the posterior end narrower and more narrowly
rounded than the anterior. The lower margin is generally gently curved. In
the female shell, which is generally more inflated, the outline is more subelliptical,
and the posterior end more broadly rounded than in the male. This is brought

RhoacFs Unio parvus (1899) from Beaver River, Wampum, Lawrence Co., refers probably to
this species.

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263

about by a slight swelling and prominence of the lower margin in the postbasal
region, and in consequence of this the lower margin is straighter in the middle, or
even slightly concave. The undulating character of the rays is generally more
pronounced in the female in the region of the postbasal swelling. On the average
the female shell also appears a little shorter and higher than the male shell.

L. H. D.

Size: (Males) 1. Utica, Cat. No. 61.3377 37 mm. 20 mm. 12 mm.

2. Sharpsville, Cat. No. 61.3378 32 “ 18 “ 10 “

3. Utica, Cat. No. 61.3377 30 “ 16 “ 9 “

4. Rosston, Cat. No. 61.2933 27 “ 15 “ 9 “

(Females) 5. Walnut Bend, Cat. No. 61.3376 31 “ 18 “ 11 “

6. Russell, Cat. No. 61.3375 29 “ 19 “ 12 “

7. Sharpsville, Cat. No. 61.3378 28 “ 17 “ 11 “

8. Russell, Cat. No. 61.3375 27 “ 17 “ 11 “

The largest ^specimen in the Carnegie Museum is a male (without locality)
40 mm. long.

Soft parts (See Ortmann, 1912, p, 339). Glochidia hitherto unknown, but I
have seen them in specimens collected in West Virginia. They are of the usual
shape, subspatulate, higher than long, but of rather small size. L. 0.17, H. 0.20 mm.

Breeding season: Gravid females were found on May 13, 1911; May 23, 1911;
May 23, 1912. In every case glochidia were present. This agrees with the
assumption that the species is hradytictic. Nothing is known about the beginning
of the breeding season.

Remarks: This species, and Toxolasma parvum, are the smallest in our fauna,
and could not be possibly confounded with any others. The differences of these
two have been pointed out under Toxolasma parvum. Eurynia fabalis can be
easily recognized by the combination of a small shell with comparatively heavy
hinge teeth, and in this* it also differs from the young of other species, for instance
E. iris. It also has a certain resemblance to young Elliptio dilatatus, but the
latter is generally longer and thinner, has less heavy teeth, more or less deeply
colored nacre, and not the peculiar rays seen in E. fabalis.

I have only one specimen from Lake Erie (in Michigan) : it is very small (13
mm. long), and has a rather bright green color. I can not tell whether there are
^ any striking differences from the normal type.

Localities in Pennsylvania represented in the Carnegie Museum:

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Mahoning River, Mahoningtown, Coverts and Edinburg, Lawi’ence Co.

Shenango River, Sharpsville, Mercer Co.

264

MEMOIRS OF THE CARNEGIE MUSEUM.

Pymatuning Creek, Pyiuatuning Township, Mercer Co.

Allegheny River, Walnut Bend, Venango Co.

Crooked Creek, Rosston, Armstrong Co.

French Creek, Utica, Venango Co.; Meadville, Crawford Co.

Connewango River, Russell, Warren Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Lake Erie, La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).

Sandusky River, Upper Sandusky, Wyandot Co., Ohio (C. Goodrich).

Swan Creek, Toledo, Lucas Co., Ohio (C. Goodrich).

IMaumee River, Fort Wayne, Allen Co., Indiana (C. Goodrich).

Ohio-drainage:

Tuscarawas River, Ohio (Holland collection).

Ohio Canal, Columbus, Franklin Co., Ohio (Smith collection).

West Fork River, Lynch Mines, Harrison Co.; Lightburn and Weston, Lewis Co., West Virginia.

Elk River, Clay, Clay Co., West Virginia. <

Tennessee-drainage :

Holston River, Mascot, Knox Co.; Noeton, Grainger Co.; Church Hill, Hawkins Co., Tennessee.

South Fork Holston River, Pactolus, Sullivan Co., Tennessee.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia.

Clinch River, Clinch River Station, Claiborne Co., Tennessee; Speers Ferry, Scott Co., Virginia; St.

Paul, Wise Co., Virginia; Cleveland, Russell Co., Virginia.

Powell River, Combs, Claiborne Co., Tennessee.

Distribution and Ecology in Pennsylvania (See fig. 27) : This is a rather rare
and local shell in western Pennsylvania, and has been found chiefly in small streams
in small numbers. It has never turned up in the Monongahela-drainage in our
state, but it is in the headwaters in West Virginia (West Fork River). Probably
the species is more abundant than the records show, but is often overlooked, or
not hunted for at the proper places. I found it in and near riffles, generally in
patches of Dianthera americana, or among other water weeds {Heteranthera, in
the iipiier Allegheny, or Potamogeton, Vallisneria, etc.). According to observa-
tions made in West Virginia it distinctly prefers these plants, in riffles, and is
deeply buried in the sand and gravel bound together by their roots and rhizomes.
By pulling up the plants it sometimes was brought to light in goodly numbers.

General distribution: Type locality, Ohio (Lea). This species belongs to the
Ohio-drainage, and from this it has crossed over into the lake-drainage in south-
eastern Michigan and northern Ohio (Conrad, 1836; Marshall, 1895; Sterki,
1907a; Walker, 1898). It is also found in the western end of Lake Erie (Walker,
1913, p. 21). It has been reported from western New York, but the only known

ORTMANN; monograph of the naiades op PENNSYLVANIA.

265

locality is Chautauqua Lake (Marshall), and this undoubtedly has been reached
from the upper Allegheny. In Indiana (Call, 1896a, 1900) it is in the Ohio and
Wabash basins, and in some of the lakes in the northern part of this drainage,
Tippecanoe (Wilson & Clark, 1912a) Winona and Pike (Norris, 1902; Headlee,
1906), and here enters the Lake Michigan-drainage, St. Joseph basin (Call, 1900).
From Illinois it has been reported from the Wabash River (Baker, 1906).

It is in the upper Monongahela and Elk River in West Virginia. Records from
Kentucky are lacking and Wilson & Clark (1914) do not report it from the Cumber-
land, but it is known from the Tennessee-drainage in Tennessee, Duck River,
Columbia, Maury Co. (Marshall, 1895), and is rather abundant in the headwaters
in East Tennessee, going up here to Virginia.

It is unknown to the south and the west of this range.

Little is known as to its ecology. Headlee (1906) gives some particulars with
regard to its occurrence in lakes.

Eurynia (Micromya) iris (Lea) (1830).

Lampsilis iris (Lea) Simpson, 1914, p. 113.

Plate XVI, figs. 6, 7.

Records frorn Pennsylvania :

Harn, 1891 (western Pennsylvania).

Simpson, 1900, p. 553 (Beaver River, Pennsylvania) (as L. fatuus)y‘‘^

Ortmann, 19095, p. 191.

Characters of the shell: Shell of medium size, moderately thick anteriorly,
rather thin posteriorly. Outline subelliptical or subovate, moderately elongated,
about twice as long as high. Anterior and posterior ends rounded. Upper and
lower margins gently convex, the lower sometimes nearly straight. Beaks not
much elevated. Beak-sculpture consisting of four to six fine, but distinct, bars,
the first subconcentric, the others distinctly double-looped. Sometimes the later
bars become irregular. Valves convex, flattened upon the sides. No distinct
posterior ridge.

Epidermis yellowish to light green, with dark green rays. The rays are more
or less distinct, sharply or poorly defined, rather narrow, straight, and mostly
not interrupted. Often there are concentric bands of color, indicating, in part,
the growth-rests.

Hinge well-developed. Pseudocardinals two in left, one in right valve, tri-
angular, a little compressed and crenulated, not heavy. Interdentum absent.

A specimen so labeled by Simpson is in the Carnegie Museum.

266

MEMOIRS OF THE CARNEGIE MUSEUM.

Laterals long, thin and narrow. Beak-cavity shallow. Dorsal muscle-scars in
beak-cavity. Anterior adductor-scars distinct and well impressed, those posterior
faint and indistinct. Nacre silvery white, highly iridescent posteriorly.

Sexual differences present in the shell, but not always very striking. The
male shell has the posterior end, more or less attenuated, narrower than the anterior
end, and the lower margin has a rather regular curve. In the female shell the
lower margin is a little expanded in the postbasal region, so that its middle part
is nearly straight, and its posterior part curves up more strongly. In consequence
of this, the posterior end of the shell is more broadly rounded, and more nearly
resembles the anterior end. There are cases, in which the female character is
very slightly developed, and others, where males do not show very well the posterior
tapering of the shell. As a rule, however, the sexes are rather easily distinguishable.
Female shells seem to be smaller on the average than male shells.

L. H. D.

Size: I. Edinburg, Cat. No. 61.3531 (probably cf) 92 mm. 47 mm. 32 mm.^^^

2. New Galilee, Cat. No. 61.1797 (sex ?) 71 “ 34 “ 22 “

3. New Galilee, Cat. No. 61.3265 (d") 68 “ 36 “ 23 “

4. Pulaski, Cat. No. 61.4830 (cf) 55 “ 27 “ 18 “

5. New Galilee, Cat. No. 61.3265 (9 ) 49 “ 26 “ 17 “

6. Waterford, Cat. No. 61.4096 (9 gravid) 43 “ 24 “ 13 “

Soft parts (See Ortmann, 1912, p. 341, fig. 23). Glochidia (See Ortmann,
1912, p. 342). They are identical with those of the var. novi-ehoraci according to
Surber (1912, PI. 3, fig. 46). Surber gives the measurements: 0.240 X 0.300,
while I gave: 0.22 X 0.28 mm.

Breeding season: The following records are at hand: Sept. 14, 1909; Sept.
18, 1917; and May 11, 1907; May 13, 1910; May 13, 1911; May 23, 1911; May
24, 1911; July 30, 1914. All my specimens had glochidia. Surber (1912, p. 7)
found glochidia in September, and Wilson & Clark (1912) report this species as
gravid (in Kankakee River) on July 28, and Aug. 3. All this speaks for a brady-
tictic form.

Remarks: This is a shell well characterized by size, shape, and color. It can
not be easily confounded with any other Pennsylvanian species. However, young
specimens somewhat resemble E. f abatis, but are distinguishable at a glance by the
relatively thinner shell, and weaker hinge teeth. The extreme brilliancy of the
nacre in the posterior section of the shell is also characteristic of this species.
The latter character distinguishes E. iris also from young Lampsilis luteola, which
resembles it to some extent. In addition L. luteola has more distinctly defined

A giant, and apparently an exceptional case.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

267

rays (if rays occur) and no concentric bands indicating growth-rests, by which
fact the young of L. luteola may be recognized as being juvenile. Old specimens
of L. luteola are much larger.

There is a great deal of variability in the character of the rays of E. iris, and
below I shall discuss a northern race (novi-eboraci) , the chief difference of which
consists in the color-pattern. Otherwise our specimens of E. iris are very uniform
in their characters.

Localities in Pennsylvania represented in the Carnegie Museum:

Little Beaver Creek, Cannelton(Miss Vera White) and New Galilee, Beaver Co.; Enon Valley, Lawrence Co.
Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith)

Slipperyrock Creek, Wurtemberg, Lawrence Co.

Mahoniirg River, Mahoningtown and Edinburg, Lawrence Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Jamestown, Mercer Co.

Neshannock Creek, Eastbrook and Volant, Lawrence Co.; Leesburg, Mercer Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

Buffalo Creek, Harbison, Butler Co.

Crooked Creek, Rosston, Armstrong Co.

Sandy Creek, Sandy Lake, Mercer Co.

French Creek, Cochranton, Crawford Co.

Leboeuf Creek, Waterford, Erie Co.

Dunkard Creek, Wiley, Greene Co.

Cheat River, Cheat Haven, Fayette Co.

Other localities represented in the Carnegie Museum:

West Fork River, Lynch Mines, Harrison Co.; Lightburn and Weston, Lewis Co., West Virginia.

Little Kanawha River, Burnsville, Braxton Co., West Virginia.

North Fork Hughes River, Cornwallis, Ritchie Co., West Virginia.

Elk River, Shelton, Clay Co.; and Sutton, Braxton Co., West Virginia.

Little Coal River, Boone Co., West Virginia (Hartman collection).

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

Distribution and Ecology in Pennsylvania (See fig. 27): Like the preceding
species {E. fabalis) E. iris is restricted to the Beaver-drainage and certain tribu-
taries of the Allegheny and Monongahela, and has been found in addition in Little
Beaver Creek. It is also present in the upper Monongahela in West Virginia.
It has never been found in the large rivers. It seems to prefer localities like those
frequented by E. fabalis (Dianthera-psitches) , and is altogether a rare shell.

General distribution: Type locality, Ohio (Lea).

The exact range of this species is hard to determine, since it has often been
confounded with its var. novi-eboraci, and because there are kindred forms in the

One specimen determined by Simpson as L. fatuus (Lea).

268

MEMOIRS OF THE CARNEGIE MUSEUM.

Tenncssee-drainage, which are not well distinguished from it, e. g., E. nebulosa
(Conrad). Whether it occurs in the state of New York is doubtful, but it might
be found in the upper Allegheny-drainage. As the material in the Carnegie
Museum shows, it exists in the tributaries of the Ohio in West Virginia. In Ohio
it is certainly present, but Sterki (1907a) does not separate the two forms. The
form from Lake Erie undoubtedly belongs to novi-ehoraci. It ranges practically
all over Indiana (Call, 1896a and 1900), and here crosses over into the lake-drainage
according to Call. Nevertheless it is quite probable that in northern Indiana its
place is taken by the variety; at all events, specimens from Winona Lake in the
Carnegie Museum belong to the latter. In Illinois it is reported from the northern
half of the state (Baker, 1906), but the common form of the Chicago area is, ac-
cording to Baker’s description (1898) without doubt the var. novi-ehoraci. Still
in this region it may pass into the typical form.

From the foregoing it apiiears that the typical E. iris belongs to the Ohio-
drainage in western Pennsylvania, West Virginia, Ohio, Indiana, and possibly
Illinois. It may cross over into the lake-drainage, but it seems to be generally
represented there by the var. novi-ehoraci. Farther to the south it seems to be
absent. It has indeed been recorded from the Tennessee-drainage, and closely
allied forms certainly occur there, but these require further study. A few other
records, such as Wisconsin (Simpson), Louisiana and Texas (Baker, 1898) are
extremely doubtful.

Strangely enough, it turns up again in the Ozark region in southern Missouri.
Utterback (1916) reports it from the basins of White and Black Rivers, and speci-
mens in the Carnegie IMuseum (James River) are absolutely indistinguishable
from Pennsylvanian specimens.

Eurynia (Micromya) iris novi-eboraci (Lea) (1838).

Simpson, 1914, p. 116, makes this a synonym of Lamj)silis iris (Lea).

Plate XVI, figs. 8, 9.

Records from Pennsylvania wanting hitherto.

Characters of variety: This form differs from the typical E. iris by the color-
pattern. The rays are not fine and more or less continuous, but rather broad,
distinct, and more or less interrupted and dissolved into dark, almost black, squarish
spots. These s])ots are generally arranged in concentric bands, so that the color-
pattern becomes very attractive.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

269

L. H. D.

Size: 1. West Springfield, Cat. No. 61.4091 (c?’) 65 mm. 35 mm. 23 mm.

2. do. “ “ do. (9) 51 “ 29 “ 18 “

3. do. “ “ do. (ci’) 48 “ 28 “ 17 “

4. do. “ “ do. (9 gravid)... 44 “ 24 “ 13 “

5. Erie, Cat. No. 61.4832 (d") 56 “ 32 “ 21 “

6. do. Cat. No. 61.4831 ( 9 ) 56 “ 29 “ 20 “

7. do. Cat. No. 61.4833 (d") 54 “ 28 “ 17 “

8. do. Cat. No. 61.4093 (9) 45 “ 25 “ 16 “

Soft 'parts identical with those of typical E. iris (See Ortmann, 1912, p. 341).
Glochidia (See Lea, Obs. VI, 1858, PL 5, fig. 14; Ortmann, 19115, PI. 89, fig. 20
as of E. iris).

Breeding season: May 23, 1909, is the only date, at which I found gravid
females. They all had glochidia.

Remarks: The color-pattern is the chief diagnostic character of this form.
In addition the ground-color of the epidermis is generally lighter than in the typical
E. iris, but this is not constant. The lake-form differs from that of Conneaut
Creek by often (but not always) having the epidermis chestnut-brown especially
toward the beaks.

Intergrades seem to be present. Baker (1898a) mentioned that the color
varies in the Chicago area, and I have transitional specimens from Winona Lake.
Even in Lea’s types the typical color-pattern is not very well developed.

It should be mentioned, that in shells from the Cumberland and Tennessee
drainages belonging to this type, the broken and spotted rays turn up frequently.
This will be discussed elsewhere. The geographical relations of these forms which
are generally referred to E. nebulosa Conrad, are still obscure.

Localities in Penns'ylvania represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, Erie, Erie Co.

Conneaut Creek, West Springfield, Erie Co.

Other localities represented in the Carnegie Museum.

Mohawk River, New York (Smith collection).

Genesee River, Rochester, Monroe Co., New York (R. H. Santens).

Black Creek, Chili, Monroe Co., New York (R. H. Santens).

Lake Erie, Port Rowan, Norfolk Co., Ontario, Canada (C. Goodrich).

Grand River, Cayuga, Haldimand Co., Ontario, Canada (C. Goodrich).

Sandusky River, Upper Sandusky, Wyandot Co., Ohio (C. Goodrich).

Swan Creek, Toledo, Lucas Co., Ohio (C. Goodrich).

Raisin River, Grape P. O., Monroe Co.; Adrian and Tecumseh, Lenawee Co., Michigan (C. Goodrich).
Winona Lake, Kosciusko Co., Indiana (E. B. Williamson).

Sheyenne River, Argusville, Cass Co., North Dakota (S. M. Edwards).

270

MEMOIRS OF THE CARNEGIE MUSEUM.

Distribution and Ecology (See fig. 27) : Type locality, Oak Orchard Creek,
Orleans Co., New York (Lea).

This form apparently is a northern race of E. iris, and is found along the
northern edge of the range of the latter in the lake-drainage. Most of the localities
known are in the St. Lawrence-drainage in New York (Marshall, 1895, but quoted
generally as iris). It is known from the Erie Canal, in Onondaga Co., and from
Oneida Co., and by this route it may have crossed over into the Mohawk River
(Carnegie Museum).

Our records show that it is in Lake Erie and some of its tributaries in Canada,
Pennsylvania, Ohio, and Michigan, and very likely the records from this region
given by Sterki (1907a) and Walker (1913) are to be referred to this variety. In
Michigan it is generally distributed, and since Walker (1898) calls it novi-eboraci
there is no doubt about it.

Farther west its presence is doubtful. It occurs in Winona Lake (Wabash-
drainage !), and Baker’s figures make it probable that it is found also in the Chicago
area, but particulars are lacking.

From Illinois the range certainly extends farther northwestward, as is shown
by our specimens from the drainage of the Red River of the North in North Dakota.
This locality is altogether new, for even the typical form has been only doubtfully
reported from Wisconsin, without exact locality, and has never been found beyond
this state.

I found this variety only once in a creek (Conneaut Creek), and here it was
abundant in riffles in fine gravel and sand. In Lake Erie it lives in from one to two
feet of water on sandy bottom, and often among a scanty growth of rushes {Juncus
americanus) .

Subgenus Eurynia Ortmann (1912).

Ortmann, 1912, p. 338.^^^

Type Unio recta Lamarck.

Two species exist in Pennsylvania.

Key to the Species of the Subgenus Eurynia.

Ri. Shell of medium size, rather thin, more or less compressed, with a rather distinct, but rounded posterior

ridge. Color of epidermis olive-green to olive-brown E. (E.) nasuta.

a 2. Shell large, thick, rather swollen, with the posterior ridge obliterated. Color of epidermis dark

green to black, rarely brownish E. (E.) recta.

Simpson’s (1914, p. 60) conception of the “ subgenus ” Eurynia differs entirely from ours.

ORTMANN: monograph of the naiades op PENNSYLVANIA.

271

Eurynia (Eurynia) nasuta (Say) (1817).^^®

Lampsilis nasuta (Say) Simpson, 1914, p. 97.

Plate XVI, figs. 10, 11.

Records from Pennsylvania:

Say, 1817 (Delaware and Schuylkill Rivers).

Gabb, 1861 (Schuylkill River and League Island, Philadelphia; “ Little Perkionien Creek
Hartman & Michener, 1874 (Schuylkill River, Chester Co.).

Marshall, 1895 (Philadelphia).

Schick, 1895 (Delaware and Schuylkill Rivers, Philadelphia).

Ortmann, 19095, p. 202, 205.

Characters of the shell: Shell of medium size and medium thickness. Outline
subelliptical or sublanceolate, elongated, distinctly over twice as long as high.
Anterior margin rounded. Lower margin more or less curved. Upper margin
straight or gently curved, forming (at least when young) a distinct angle with the
posterior margin, which slopes down obliquely, joining the lower margin in a dis-
tinct, but rounded, posterior angle or point. Thus the shell is distinctly attenuated
and pointed behind, although the point itself is narrowly rounded. Beaks low,
hardly elevated above the hinge-line. Beak-sculpture (Marshall, 1890, fig. 5)
consisting of five to seven fine bars, the first one or two subconcentric, the following
double-looped, with a distinct re-entering angle or sinus back of the middle. An-
terior loop broadly rounded, posterior loop narrow, somewhat angular upon the
posterior ridge, and indistinct upon the posterior slope. Valves moderately
convex, flattened upon the sides. A posterior ridge is present; it is most distinct
near the beaks, and angular, farther downwards it becomes broader, is more
rounded and indistinct. Posterior slope slightly concave near the beaks, becoming
flatter or even somewhat convex toward the posterior end of the shell. In young
shells it is compressed and elevated toward the posterior angle of the upper margin.

Epidermis dark olive-green or brown, with or without rays. The latter are
poorly developed, dark green, straight, narrow or somewhat wider, and generally
visible only (if at all) in the posterior section of the shell, just in front of the pos-
terior ridge, and upon the posterior slope. In younger specimeps the rays may
extend over the whole surface. In other cases, they are entirely absent. Some-
times there are concentric light and dark bands, the latter marking the growth-
rests.

176 Not 1816.

177 Perkiomen Creek is a tributary of the Schuylkill in Montgomery County. There is no “ Little
Perkiomen Creek ” to my knowledge.

272

MEMOIRS OF THE CARNEGIE MUSEUM.

Hinge well-developed. Pseudocardinals one or two in left, one or two in
right valve, not very large and not heavy, compressed, siibtriangular, somewhat
crenulated. Laterals long and thin. No interdentum. Beak-cavity shallow.
Dorsal muscle-scars in beak-cavity. Adductor-scars distinct, and moderately
impressed anteriorly, less distinct posteriorly. Nacre silvery white, often cream-
color or salmon toward the beak-cavity, highly iridescent posteriorly.

Sexual differences of the shell well marked. In the male the shell tapers
uniformly behind into the ])osterior point, and the lower margin has a rather
uniform curve. In the female the lower margin is distinctly produced in the
postbasal jiortion, there forming a broad, rounded projection, behind' which the
lower margin slopes up more suddenly, and is at this point straight or even slightly

concave.

L. H. D.

Size: 1. Yardley, Cat. No. 61. .3525 (largest cf’) 77 mm. 35 mm. 19 mm.

2. Erie, Cat. No. 61.4088 (cf) 102 “ 44 “ 30 “

3. do. Cat. No. 61.4837 (9) 93 “ 45 “ 27 “

4. do. Cat. No. 61.4083 (d^) 89 “ 42 “ 24 “

5. do. Cat. No. 61.4088 (9 ) 71 “ 33 “ 17 “

6. do. “ “ do. (9) 58 “ 28 “ 15 “

Soft parts (See Ortmann, 1912, p. 343). Glochidia (See Lea, Obs. XIII, 1874,
PI. 21, fig. 2; Ortmann, 1912, PI. 20, fig. 8).

Breeding season: Conner (1907) mentions this species as breeding all the year
round. From the eastern jiart of the state I have only two records for gravid
females: Sept. 15, 1905 and May 10, 1909, in both cases with glochidia. From
Lake Erie I have the following records: end of August, 1909 (eggs), and May 21,
1909; May 22, 1909; May 24, 1909; June 2, 1908; June 3, 1908; July 7, 1910.
The dates in early summer furnished glochidia, and on the two last ones, dis-
charging females were found. On and after July 8 numerous females were col-
lected, but none were gravid, so that in Lake Erie, as I have shown (1912, p. 343),
a distinct interim exists in July and at the beginning of August.

liemarks: This is a species quite easily recognized by its lance-head outline,
dull olive-green color, with only indistinct rays, and light-colored nacre. How-
ever, it resembles, principally in the male sex certain other species, for instance
Elliptio cupreus (Rafinesque) and E. fisherianus (Lea), and indeed, specimens of
the latter have been confounded by Conrad with it. E. cupreus and E. fisherianus
generally have a differently colored nacre (coppery or purple), and differ in the

In old males, there is sometimes a short concavity of the lower margin immediately in front of
the posterior end. This is due to a downward deflection of the posterior end, such as is often observed in
old specimens of other species.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

273

hinge-teeth, which are more stumpy. The females of these two are entirely
different from those of E. nasuta, and have never the peculiar expansion of the
lower margin.

There is not much variability in E. nasuta. The specimens may be a little
longer or shorter in outline, and the color of the epidermis may b'e more greenish
or more brownish. In this connection it should be mentioned that specimens
from Lake Erie frequently differ from eastern specimens by their rusty brown

• Eurynia nasuta (eastern range).

color, especially towards the beaks. This is the effect of the lake environment,
which is observed in other species from the lake. But this color is not always
present. As usual, specimens from Lake Erie often have very regular growth-
rests. In aU other characters, the lake-form is absolutely like that of eastern
Pennsylvania.

Localities in Pennsylvania represented in the Carnegie Museum:

Lake Erie, Miles Grove, and Presque Isle Bay, also beach-pools of Presque Isle, Erie, Erie Co.

Delaware River, Penns Manor and Yardley, Bucks Co.

Schuylkill Canal, Manayunk, Philadelphia Co.

Other localities represented in the Carnegie Museum:

Douglas Lake, Cheboygan Co., Michigan (H. B. Baker).

Otter Creek, Monroe Co., Michigan (C. Goodrich).

Ottawa River, Toledo, Lucas Co., Ohio (C. Goodrich).

Lake Erie, La Plaisance Bay, Monroe Co., Michigan (C. Goodrich); Sandusky Bay, Cedar Point, Erie
Co., Ohio (0. E. Jennings; C. Brookover); Port Rowan, Norfolk Co., Ontario, Canada (C. Good-
rich); Crystal Beach, Welland Co., Ontario, Canada (F. Behrle).

Delaware-Raritan Canal, Princeton, Mercer Co., New Jersey.

Delaware River, Fish House, Camden Co., New Jersey.

274

MEMOIRS OF THE CARNEGIE MUSEUM.

Localities represented in the Philadelphia Academy:

Delaware River, Kaighn’s Point, Camden, Camden Co., New Jersey (John Ford); Delanco (H. A.
Pilsbry), Burlington Island (H. W. Fowler), and Florence (H. W. Fowler), Burlington Co., New
Jersey.

Swartswood Lake, Sussex Co., New Jersey (H. A. Pilsbry & S. N. Rhoads), drains to Delaware River.
Cohansey Creek, Bridgeton, Cumberland Co., New Jersey (S. N. Rhoads) coastal plain of southern New
Jersey.

Potomac River, Washington, D. C. (John Ford); Alexandria, Fairfax Co., Virginia (G. W. Tryon, Jr.).

Distribution and Ecology in Pennsylvania (See figs. 28 & 29) : In Pennsylvania
this species has two ranges: one in the extreme eastern end of the state, and the
other in the northwest, in Lake Erie. Its metropolis in the east is found in the
tidewaters of the Delaware River; farther up (above Trenton, New Jersey) it is

• Eurynia nasuta (western range).

+ Eurynia recta.

rare. It also has been reported from the Schuylkill, probably its lower part,
as far up as Chester Co. The record from Perkiomen Creek in Montgomery Co.
is doubtful (See above). In addition I found a single individual in the Schuylkill
Canal. It has never been found in any of the smaller tributaries, and no records
are at hand from anywhere in the Susquehanna or Potomac drainages in Penn-
sylvania.

I found it in large numbers in the tidewater of the Delaware opposite Phila-
delphia at Fish House. Here it inhabits the shores of the Delaware, which re-
sembles rather a lake than a river. The bottom at this point is fine sand, and the

ORTMANN: monograph of the naiades of PENNSYLVANIA.

275

water is subject to the tides. At Penn’s Manor I found it in the deep water of a
protected cove, on a bottom consisting of mud and vegetable debris. The single
specimen from the Schuylkill Canal was in deep black muck, the only one found at
Yardley lived in gravel in an eddy in a riffle.

In Lake Erie it is a very common species in Presque Isle Bay, where it is
present in all parts on sandy, gravelly, and muddy bottom, in from one to fifteen
feet of water, and it is also abundant in certain beach-pools of Presque Isle, on
sandy and sandy-muddy bottom.

Sandy bottom of great, quiet bodies of water (tidewaters, lakes and probably
also canals) seem to furnish the conditions most favorable to this species.

General distribution: Type locality, Schuylkill and Delaware Rivers, Phila-
delphia (Say).

The distribution of this species is unique: only Elliptio violaceus can be com-
pared with it, but even this is by no means identical (See under E. violaceus, p. 110,
and Walker, 1913). It is found on the Atlantic coastal plain, but does not go
far up into the rivers, at least in the southern part of its range. Although Simpson
(1900) reports it as far South as North Carolina, I have been able to find as its
most southern record only that of Conrad (1836) from James River, Virginia (the
figured specimen is from this place). Conrad also gives the Potomac river at
Washington, and this is confirmed by other writers (Dewey, 1856; Marshall,
1895) and by specimens in the Philadelphia Academy. It is not known from
anywhere in the headwaters of the Atlantic streams in Virginia and Maryland,
and I myself never found it in the upper James, the upper Rappahannock, and the
Potomac, on the Piedmont Plateau and West of the Blue Ridge. It has been
reported from the lowlands in Delaware (Rhoads, 1904). In Pennsylvania its
distribution is restricted chiefly to the lowlands, and it is absent in the upper
Susquehanna. In New Jersey it is known from the Delaware River and from the
coastal plain (Philadelphia Academy). I found it in the Delaware-Raritan Canal,
and in addition there is a locality (given above) in Sussex Co., northern New Jersey,
belonging to the upper Delaware-drainage, which is highly interesting. It is
found in New York in the Hudson River as far up as Troy and Albany (Aldrich,
1869; Marshall, 1895^'^®), and a number of localities are known from Connecticut,
Rhode Island, and Massachusetts (Linsley, 1845; Perkins, 1869; Carpenter, 1890;
Earle, 1835; Gould-Binney, 1870; Marshall, 1895; Johnson, 1915). It goes
northward to Keene, Cheshire Co., New Hampshire (Walker & Coolidge, 1908).

From the Hudson-drainage in New York, this species ranges westward through
the Mohawk basin and the Erie canal, and enters the St. Lawrence-drainage.

Marshall also gives Delaware River, but no exact locality.

276

MEMOIRS OF THE CARNEGIE MUSEUM.

Here it is found in the tributaries of Lake Ontario in New York, and in Hamilton
Bay of Lake Ontario in Canada (DeKay, 1843; Marshall, 1895), and reaches the
Niagara River and Lake Erie (Marshall). In Lake Erie it extends farther west-
wards, along the Canadian (Carnegie Museum) Pennsylvanian and Ohio shores
(Sterki, 1907a; Walker, 1913). In Ohio it is found also in tributaries of the lake
in Portage Co. (Dean, 1890, Sterki) and Ciij^ahoga River (Dean, 1890), and in
Ottawa River, Lucas Co. (Carnegie Museum). Further it is found in Michigan
over the whole state (Walker, 1898). Call (1896a) reports it also from the St.
Joseph basin in Indiana, but in 1900 (p. 458) he says that no authentic specimens
are known from this state. Farther to the south and west it seems to be replaced
by the closely allied E. suhrostrata (Say).

The western and the eastern ranges seem to be connected from Lake Erie to
the Hudson River by the Erie canal, and it is very likely that this species migvated
along this route. The direction of the migration probably was from west to
east, considering the fact that the only closely allied species, Eurynia suhrostrata,
is a species of the interior basin. Being a species partial to quiet water, it is only
natural that it followed the route of the Erie canal. In this case, however, this
migration must have been quite recent, and it must be a very late arrival on the
coastal plain, where it has spread in recent times, as far South as Virginia, remaining
restricted chiefly in the southern part of its range to the Coastal plain. This has
been discussed previously in another paper (Ortmann, 1913a, p. 378 et seq.).

Eurynia (Eurynia) recta (Lamarck) (1819).

Lampsilis recta (Lamarck) Simpson, 1914, p. 95.

Plate XVI, figs. 12, 13.

Records from Pennsylvania:

Harn, 1891 (western Pennsylvania).

Clapp, 1895 (Allegheny Co.)

Marshall, 1895 (Allegheny River, Warren Co.)

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.)

Ortmann, 19095, pp. 190 and 202.

Characters of the shell: Shell large, solid and heavy. Outline subelliptical to
sublanceolate, elongated, distinctly over twice (up to two and a half times) as
long as high. Anterior margin rounded; lower margin more or less convex.
Upper margin nearly straight, passing gradually into the descending posterior

Sterki says: “ probably also in the Ohio-drainage, at least along the divide.”

This route is more southern than that of Elliptio violaceus, where the connection does not go
through Lake Erie but is more northern.

ORTMANN: monograph of the naiades of PENNSYLVANIA,

277

margin, which joins the lower margin in a distinct, but rounded, angle, so that the
shell is more or less narrowed and pointed at the posterior end. Beaks low, only
little elevated above the hinge-line. Beak-sculpture faint and obsolete, con-
sisting of three to five indistinct bars, which are double-looped, with a sinus in
the middle; the posterior loop is open, i.e., the bars disappear upon the posterior
slope. Valves moderately convex, flattened upon the sides. Posterior ridge
practically absent, only toward the beaks is it faintly indicated. Posterior slope
gently convex, flat towards the beaks.

Epidermis glossy, dark green to brown or blackish, with indistinct rays or
without rays. Young shells are lighter green, with dark green rays of various
width, but in the posterior section of the shell the rays are generally rather wide.
In old shells, the rays become obscure, and the epidermis is often uniformly dark
brown or black. Sometimes concentric bands of dark color, marking growth-
rests, are present, but generally the growth-rests are not marked by color.

Hinge well-developed. Pseudocardinals two in left, one or two in right valve,
stumpy or slightly compressed, crenulated. No interdentum. Laterals long,
strong, rather straight. Beak-cavity shallow. Dorsal muscle-scars in beak-cavity.
Adductor-scars distinct, those anterior well impressed, those posterior less so.
Nacre white, iridescent posteriorly. Often there is more or less pink or light
purple in the beak-cavity and upon the hinge teeth; sometimes all of the nacre is
purple, but such specimens have not been found in Pennsylvania.

Sexual differences of shell well marked. In the male, the shell tapers pos-
teriorly rather uniformly to a point, and the posterior part is distinctly narrower
than the anterior. In the female, the lower margin is distinctly enlarged in the
postbasal part. In front of this widening, the lower margin is straight or slightly
concave, behind it slopes up rather suddenly to the posterior point, which, in
consequence of this, is more elevated above the base line than in the male. The
posterior section of the female shell is not distinctly narrowed, and sometimes even
slightly higher, than the anterior. However, in some females, the characteristic

shape is not very well developed.

L. H. D.

Size: (Males) 1. Aladdin, Cat. No. 61.3522 177 mm. 70 mm. 49 mm.

2. Neville Island, Cat. No. 61.706 175 “ 70 “ 51 “

3. Industry, Cat. No. 61.4077 100 “ 41 “ 27 “

4. Mosgrove, Cat. No. 61.4075 71 “ 31 “ 16 “

(Females) 5. Cooks Ferry, Cat. No. 61.3519 156 “ 67 “ 52 “

6. Industry, Cat. No. 61.4077 140 “ 60 “ 38 “

7. Industry, Cat. No. 61.3518 Ill “ 47 “ 31 “

8. Meadville, Cat. No. 61.3512 89 “ 40 “ 23 “

{Lake Erie-form) 9. Erie, Cat. No. 61.4070 (cf) 118 “ 47 “ 33 “

10. do. “ “ do. (9) 112 “ 52 “ 31 “

278

MEMOIRS OF THE CARNEGIE MUSEUM.

The species is said to grow even larger than the largest specimens listed above,
but Pennsylvanian specimens compare favorably with specimens from other
localities.

Soft parts (See Lefevre & Curtis, 1910, PL 1, fig. 5; 1912, PI. 7, fig. 6; Ort-
mann, 1912, p. 344, fig. 24). Glochidia: Lea, Obs. VI, 1858, PI. 5, fig. 11; Lefevre
& Curtis, 1910, p. 97, fig. L; Ortmann, 19116, PL 89, fig. 21; Lefevre & Curtis,
1912, p. 146, fig. L; Surber, 1912, PL 2, fig. 17. My measurements are: 0.22 X
0.28; those of Lefevre & Curtis : 0.20 X 0,24; those of Surber : 0.220 X 0.280 mm.

Breeding season: I have the following records of gravid females: Aug. 13,
1907; Aug. 29, 1908; Aug. 30, 1909; Sept. 1, 1908; Sept. 3, 1908; Sept. 5, 1908;
Sept. 6, 1908; Sept. 7, 1914; Sept. 8, 1908; Sept. 11, 1913; Sept. 12, 1913; Sept.
14, 1915; Sept. 15, 1909; Sept. 17, 1913; Sept. 17, 1915; Sept. 22, 1910; Sept.
23, 1908; Sept. 24, 1917; Oct. 24, 1907; Oct. 24, 1910; Nov. 4, 1914; Nov. 12,
1911; and then again: May 21, 1909; May 22, 1909; May 23, 1914; May 25,
1914; July 5, 1909; July 10, 1908; July 23, 1907.

Eggs have been found in August (earliest date Aug. 13). In September and
October specimens with glochidia are present, and then again in May. But all
three records in July refer to specimens with glochidia. Thus the breeding season
lasts from about the middle of August to the end of July, and may overlap with the
next, or there ma}" be a short interval at the end of July and in the first half of
August. Surber found glochidia in September, October, November, in March,
April, and May, and then again in July, which agTees well with my records. The
species is bradytictic.

Remarks: Eurynia recta generally is easily recognized by its large size, elon-
gated shape, and almost black color. However, young male specimens sometimes
resemble Elliptio dilatatus, but may be distinguished by the shining, green-black
epidermis, with indistinct, broad rays (if such are at all visible). Elliptio dilatatus
has a dull, more brownish-black epidermis, with fine rays (if such are at all visible).
The posterior end of E. recta is also more pointed, and the nacre is generally not
so deeply colored as in E. dilatatus. There are additional differences in the hinge-
teeth and the beak-sculpture. The female shell of E. recta cannot be confounded
with E. dilatatus.

Eurynia recta is not very variable. The proportion of length and height
varies slightly, and so does the color of the epidermis and of the nacre. In Lake
Erie there is a local form, which differs from the Ohio type by its smaller size, its
paler epidermis (browner, inclinding to russet), and more regular growth-lines.
But there are specimens in the lake, chiefly younger ones, which are indistinguish-

OETMANN: MONOGKAPH of the naiades of PENNSYLVANIA.

279

able from specimens from the Ohio River. Therefore I do not think it advisable
to separate the two forms by varietal names. Specimens from Maumee River
are entirely normal, i-e., like the Ohio-form.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio-drainage:

Ohio River, Shippingport, Cook’s Ferry, and Industry, Beaver Co.; Neville Island, Allegheny Co.; Edge-
worth, Allegheny Co. (G. H. Clapp).

Allegheny-drainage :

Allegheny River, Schenley, Aladdin, Godfrey, White Rock (Johnetta), Kelly, Mosgrove, and Templeton,
Armstrong Co.; Walnut Bend, Venango Co.; Tionesta and Hickory, Forest Co.; Warren, War-
ren Co.

Crooked Creek, Rosston, Armstrong Co.

French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co.
Connewango Creek, Russell, Warren Co. (Dr. R. R. Jones).

Monongahela-drainage:

Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).

Dunkard Creek, Wiley, Greene Co.

Cheat River, Cheat Haven, Fayette Co.

Lake-drainage :

Lake Erie, Presque Isle Bay, and beach-pools of Presque Isle, Erie, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Severn River, Gloucester Pool, Muskoka Co., Ontario, Canada (0. A. Peterson).

Lake Erie, Port Rowan, Norfolk Co., Ontario, Canada (C. Goodrich); Cedar Point, Erie Co., Ohio
(C. Brookover); La Plaisance Bay, Monroe Co., Michigan (C. Goodrich).

Maumee River, Waterville, Lucas Co.; and Otsego Rapids, Wood Co., Ohio (C. Goodrich).

Ohio-drainage:

Tuscarawas River, Ohio (Holland collection).

Ohio River, Congo, Hancock Co., West Virginia; Toronto, Jefferson Co., Ohio; St. Marys, Pleasants
Co., West Virginia; Parkersburg, Wood Co., West Virginia; Portland, Meigs Co., Ohio.

Cheat River, Jaco and Mont Chateau, Monongalia Co., West Virginia.

Little Kanawha River, Grantsville, Calhoun Co., West Virginia (W. F. Graham).

Elk River, Shelton and Clay, Clay Co.; Gassaway, Braxton Co., West Virginia.

Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.

It should be borne in mind that the original recta of Lamarck is from Lake Erie, and was sub-
sequently called sageri by Conrad (See Walker, 1913, p. 21). The name recta should be retained for the
lake-form, while for the form from the Ohio the varietal name latissima Rafinesque is available.

Iii addition it may be stated that dead shells have been seen, but not taken, in the Conemaugh
River, New Florence, Westmoreland Co.

280

MEMOIRS OF THE CARNEGIE MUSEUM.

Tennessee-drainage :

Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith); Concord and Knoxville, Knox Co.,
Tennessee.

French Broad River, Boyd Creek, Sevier Co., Tennessee.

Nolichucky River, Chiinns Shoals, Hambleir Co., Tennessee.

Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill, Noeton, and
Holston Station, Grainger Co., Tennessee.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee.

Clinch River, Solway, Knox Co.; Edgemoor, Clinton, and Offutt, Anderson Co.; Clinch River Station,
Claiborne Co.; Oakman, Grainger Co., Tennessee; Clinchport,. Scott Co., Virginia; St. Paul,
Wise Co., Virginia.

Powell River, Combs, Claiborne Co., Tennessee.

Mississippi-drainage and westward:

Kaskaskia River, Illinois (G. H. Clapp, donor).

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection); Moline, Rock Island Co.,
Illinois (P. E. Nordgren).

Kishwaukee River, Rockford, Winnebago Co., Illinois (P. E. Nordgren).

Meramec River, Meramec Highlands, St. Louis Co., Missouri (N. M. Grier).

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

White River, Cotter, Baxter Co., Arkansas (A. A. Hinkley).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler).

Saline River, Benton, Saline Co., Arkansas (H. E. Wheeler).

Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler).

Neosho River, Miami, Ottawa Co., Oklahoma (F. B. Isely).

Alabama-drainage :

Coosa River, Wilsonville, Shelby Co.; Riverside, St. Clair Co., Alabama (H. H, Smith).

Chattooga River, Cedar Bluff, Cherokee Co., Alabama (H. H. Smith).

Distribution and Ecology in Pennsylvania (See fig. 29) : Eurynia recta is found
in Pennsylvania in Lake Erie and in the Ohio-drainage. In the latter it distinctly
prefers the larger rivers, where it is generally abundant, and attains a huge size.
In the Allegheny it goes up to Warren County. Yet it has also entered some of
the smaller tributaries. It is found in the lower part of Crooked Creek, in French
Creek (up to Cambridge Springs), and in Connewango Creek. It also must have
gone up the Kiskiminetas, for dead shells have been seen in the Conomaugh at
New Florence. In the Monongahela-drainage it is present in the lower part of
Dunkard Creek, and it is in Cheat River, where it passes up into West Virginia.
On the other hand, its total absence in the whole Beaver-drainage is most remark-
able.

Its ecological preferences show that it is mainly a form of the big rivers. In
the Ohio it belongs chiefly to the shell-banks in the deep channel of the river with

ORTMANN: monograph of the naiades of PENNSYLVANIA.

281

gravelly bottom and strong, steady currents. In smaller streams it is also mostly
in very strong currents and in heavy gravel on riffles. It buries very deeply.
In Lake Erie it is found in the fine sand and gravel of Presque Isle Bay, in pro-
tected locations (among rushes) as well as on open, surf-beaten shores. It is also
in some of the beach-pools with quiet water and sandy-muddy bottoms.

General distribution: Type locality, Lake Erie (Lamarck).

The metropolis of E. recta is in the large rivers of the central basin. From
western Pennsylvania, it spreads all through the Ohio-drainage in Ohio (Sterki,
1907a), Indiana (Call, 1896a, 1900), in West Virginia, and eastern Kentucky.
From the rest of Kentucky records are lacking, but it is in the Cumberland (Mar-
shall, 1895; Wilson & Clark, 1914), and in the Tennessee, where it ascends in the
Clinch River to Virginia. It is common in the Mississippi-drainage in Illinois
(Baker, 1906), Iowa (Pratt, 1876; Witter, 1878; Call, 1895; Marshall, 1895;
Geiser, 1910), and northward to Wisconsin (Conrad, 1836; Cooper, 1855; Lapham,
1860), and Minnesota (Cooper, 1855; Grant, 1886; Holzinger, 1888).

In the Missouri-drainage, it is found in Missouri (Utterback, 1916), in north-
eastern Kansas (Scammon, 1906), and southeastern Nebraska (Tryon, 1868; Call,
1895). Southward it becomes less abundant, but is known from the southern
drainage in Kansas (Scammon), Arkansas (Call, 1895; Marshall, 1895; Wheeler,
1918), and Oklahoma. Its presence in Texas is doubtful.

In addition this species has crossed over into other drainages in the north
as weU as in the south. From northern Indiana and Ohio it has reached the
lake-drainage in Michigan (Walker, 1898) and Lake Erie (Walker, 1913), and has
spread down the St. Lawrence far into eastern Canada, Ottawa, Quebec, Montreal
(See Bell, 1859; Whiteaves, 1863; Call, 1885; Marshall, 1895) and has entered
some of the tributaries of the St. Lawrence in western New York (Marshall, 1895).
In Ontario it goes northwest at least to the region of Georgian Bay, Severn River
(See Carnegie Museum). Further in Minnesota it has crossed over into the
drainage of the Red River of the North, Wilkin County (Grant, 1886), and has
reached North Dakota, at Pembina (Call, 1885) and Roseau River (Dawson, 1875),
Winnipeg (Christy, 1855; and Hanham, 1899) Canada.

Finally it is found in the south in the Alabama-drainage in Alabama and
Georgia (Conrad, 1836; Lewis, 1877; Call, 1885, 1895).

As has been said, it prefers larger rivers, and within the immense area indi-
cated above, it may be missing in many of the smaller, and even medium-sized
streams. Baker (1898a) records it from lakes and large rivers, on muddy bottoms,
while Scammon (1906) says that it is found under a variety of conditions.

282

MEMOIRS OF THE CARNEGIE MUSEUM.

Genus Lampsilis Rafinesque (1820).

Ortmann, 1912, p. 345; Simpson, 1914, p. 35.

Type Unto ovatus Say.

In Pennsylvania I distinguish seven species and three varieties, which are to
be recognized as follows :

Key to the Species and Varieties of Lampsilis.
fli. Shell subelliptical, elongated, considerably over one-and-a-half times as long as high.

bi. Shell more or less swollen; in the female greatly inflated and broader behind. Epidermis more
or less smooth and shining, with the rays, when present, rather sharply defined.

Cl. Larger, greenish-yellow, with dark rays, colors bright. Growth-lines irregular and not

very distinct L. luteola.

c<i. Smaller, pale yellowish, or pale brownish, rays not so dark, colors duller. Growth-lines

generally distinct and regular L. luteola rosacea.

ho. Shell more or less compressed, in the female not so greatly inflated and broadened behind. Epi-
dermis rough, and not shining, with the rays less sharply defined L. radiata.

a 2. Shell suboval or subelliptical, not elongated, rarely about one and a half times as long as high, gen-
erally shorter.

bi. Shell moderately thick, or even comparatively thin. Color of epidermis yellowish, greenish,
or grayish olive, or dull brownish. Rays variable, but, when present, generally sharply
defined.

Cl. Epidermis yellowish or greenish olive, more or less shining (dull only in old shells). Rays,
when present, more or less sharply defined.

di. Epidermis more or less greenish, more or less shining; when yellowish, the rays cover
most of the shell.

Cl. Rays absent or simple, broader or narrower, not much crowded. Shell rather
large.

/i. Shell larger, colors brighter, rays more sharply marked, when present, and
rather dark.

gi. A sharp posterior ridge present, and posterior slope truncate. . . .L. ovata.
g2. No sharp posterior ridge, posterior slope not truncate.

L. ovata ventricosa.

/2. Shell smaller, colors duller and paler, rays less distinct and not very dark.

L. ovata canadensis.

62. Rays always present, fine, rarely somewhat broad, much crowded, and more or

less interrupted or wavy. Shell of smaller size L. fasciola.

di. Epidermis bright yellow (wax-yellow), sometimes inclining to reddish brown, very
bright and glossy. Rays, if present, very distinct, but generally restricted to the

posterior part of the shell L. cariosa.

C2. Epidermis dull in color, grayish olive, not very shining. Rays ill-defined L. ochracea.

62- Shell very tliick and heavy. Color of epidermis more or less reddish-yellow or brown. Rays
indistinct L. orbiculata.

OETMANN: monograph of the naiades of PENNSYLVANIA.

283

Lampsilis luteola (Lamarck) (1819).

Lampsilis luteola (Lamarck) Simpson, 1914, p. 60.

Plate XVII, figs. 1, 2.

Records from Pennsylvania :

Clapp, 1895 (Allegheny Co.)

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co.; Beaver, River, Wampum, Lawrence Co.)
Ortmann, 19096, p. 190, 202.

Characters of the shell: Shell from medium size to rather large, moderately
thick. Outline subelliptical or subovate, moderately elongated, distinctly over
one-and-a-half times as long as high, but less than twice as long as high. Anterior
margin rounded. Lower margin more or less regularly curved. Upper margin
nearly straight, forming with the posterior margin a blunt angle, when young, or
passing gradually into it, when old. Posterior margin meeting the lower margin
in a very blunt angle, so that the posterior end of the shell is rather rounded off.
Beaks low, located in the anterior portion of the shell. Beak-sculpture (see Mar-
shall, 1890, fig. 3) fine, consisting of six to ten double-looped bars, with a distinct
re-entering angle in the middle. The posterior loop is slightly angular, and in-
distinct upon the posterior slope. Sometimes the first bar appears subconcentric.
Valves more or less swollen, rarely subcompressed, but flattened upon the sides.
Posterior ridge indistinct.

Epidermis yellowish to light greenish, or light brownish, shining and smooth.
In old shells it is darker, but not blackish. Rays are generally present. They
are dark green to blackish, quite distinct, straight, narrow or broad, covering
more or less of the surface. Posterior slope often darker in color, and less shining.
Growth-rests more or less distinct, but irregular.

Hinge well-developed. Pseudocardinals two in left, one or two in right valve,
tooth-like, compressed or stumpy, crenulated, very variable. Interdentum absent.
Lateral teeth long, but rather thin, gently curved. Beak-cavity shallow. Dorsal
muscle-scars in beak-cavity. Adductor-scars distinct and well impressed, chiefly
the anterior one. Nacre milky-white, silvery, somewhat iridescent behind.

Sexual differences very strongly marked. While the male shell uniformly
tapers backward to the bluntly pointed end, having the lower margin rather evenly
curved, the female develops a considerable swelling and expansion of the lower
margin in the postbasal region, so that the anterior part of the lower margin is
nearly straight (or even slightly concave), and the posterior section of it ascends
very suddenly to the posterior end. This produces a considerable widening of the
posterior end of the shell, which often appears truncated. In old females this
feature is so prominent, that the shell actually seems to be distorted.

284

MEMOIRS OF THE CARNEGIE MUSEUM.

L. H. D.

Size: (Males) 1. Linesville, Cat. No. 61.4027 142 mm. 78 mm. 60 mm.

2. Jamestown, Cat. No. 61.3453 125 “ 69 “ 54 “

3. Rosston, Cat. No. 61.4036 78 “ 45 “ 23 “

(Females) 4. Linesville, Cat. No. 61.4027 (gravid) .. Ill “ 73 “ 60 “

5. Wampum, Cat. No. 61.2873 101 “ 68 “ 41 “

6. Shenango, Cat. No. 61.4100 (gravid) . . 71 “ 46 “ 26 “

The maximum length given by Baker (1898a) is 116 mm., by Scammon
(1906), 136 mm. Thus it is seen that the Pennsylvanian specimens attain an
extreme size.

Soft parts (See Ortmann, 1912, p. 348). Glochidia (See Lea, Obs. VI, 1856,
PI. 5, fig. 10; Surber, 1912, PI. 2, fig. 15). My measurements are: 0.23 X 0.28,
while Surber gives: 0.250 X 0.290 mm. But Surber remarks (1912, p. 4) that
at certain localities, the glochidia are uniformly smaller than at others.

Breeding season: My dates for gravid females are very numerous, and cover
the periods from August 3 to October 27, and from April 22 to July 19. The

• Lampsilis luteola.
■ Do. var. rosacea.

species is bradytictic, but found gravid nearly all the year round, except in July,
where there seems to be a short interim. Although gravid- (and discharging)
females have been found in July, they are rare in this condition in that month.

Remarks: A very variable species, but recognized by its subelongated shape,
the light color of the epidermis, which is shining, and has generally weU-marked

ORTMANN: monograph of the naiades of PENNSYLVANIA,

285

rays. Young specimens of Actinonaias ligamentina sometimes resemble this species,
but their shape is more broadly elliptical. Large individuals of Eurynia iris also
might be mistaken for L. luteola, but they have a more elongated shape, more
iridescent nacre, and besides the character of the rays is different.

In old specimens the epidermis is often more or less discolored, and the smooth-
ness disappears, but specimens with dark (brown or black) epidermis are very
rare. The typical inflation of the valves is best developed in old individuals,
while young ones are generally more or less compressed.

Several forms (or species) have been distinguished. The variety rosacea will
be discussed below. But there are other local forms, which, however, are not
constant enough to deserve special mention, and most of them are not found in
our state.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River and Ohio-drainage:

Ohio River, Cooks Ferry and Industry, Beaver Co.; Coraopolis (S. N. Rhoads) and Neville Island,
Allegheny Co.

Buffalo Creek, Acheson, Wasliington Co.

Little Beaver Creek, Cannelton (Miss Vera White & H. H. Smith), Darlington, and New Galilee, Beaver
Co.; Enon Valley, Lawrence Co.

Raccoon Creek, Raccoon Township, and New Sheffield, Beaver Co.; Bavington, Washington Co.
Chartiers Creek, Carnegie, Allegheny Co. (D. A. Atkinson & J. L. Graf).

Little Chartiers Creek, Morganza, Washington Co.

Beaver-drainage :

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Zelienople and Harmony, Butler Co.

Slipperyrock Creek, Wurtemberg and Rose Point, Lawi’ence Co.

Wolf Creek, Grove City, Mercer Co.

Brush Creek, Celia, Beaver Co.

Little Connoquenessing Creek, Harmony, Butler Co.

Glade Run, Zeno, Butler Co.

Thorn Creek, McBride, Butler Co.

Bonnie Brook, East Butler, Butler Co.

Mahoning River, Mahoningtown, Coverts, Edinburg, and Hillsville, Lawrence Co.

Neshannock Creek, Volant, Lawi’ence Co.; Leesburg, Mercer Co.

Shenango River, Harbor Bridge and Pulaski, I awrence Co.; Sharpsville, Clarksville, Shenango, and
Jamestown, Mercer Co.; Linesville, Crawford Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

Little Shenango River, Greenville, Mercer Co.

Randolph Run, Hartstown, Crawford Co.

Padan Creek, Linesville, Crawford Co. (0. E. Jennings).

286

MEMOIRS OF THE CARNEGIE MUSEUM.

Allegheny-drainage:

Allegheny River, Johnetta, Kelly, Mosgrove, and Templeton, Armstrong Co.; Larabee, McKean Co.
(Dennis Daily).

Crooked Creek, Rosston and South Bend, Armstrong Co.; Creekside, Indiana Co.

Cowanshannock Creek, Rural Valley, Armstrong Co. (N. M. Grier). .

Little Mahoning Creek, Goodville, Indiana Co.

French Creek, Utica, Venango Co.; Cochranton, Meadville, and Cambridge Springs, Crawford Co.
Conneaut Outlet, Conneautlake, Crawford Co.

Conneaut Lake, Cra\vford Co.

Cussewago Creek, Mosiertown, Crawford Co. (H. & L. Ellsworth).

Conneauttee Creek, Edinboro, Erie Co.

Leboeuf Creek, Waterford, Erie Co.

Connewango Creek, Russell, Warren Co.

Potato Creek, Smethport, McKean Co. (P. E. Nordgren).

Monongahela-drainage:

Monongahela River, Elizabeth, Allegheny Co. (D. A. Atkinson).

Ten Mile Creek, Clarksville, Greene Co.; Amity, Washington Co.

South Fork Ten Mile Creek, Clarksville and Waynesburg, Greene Co.

Dunkard Creek, Wiley and Mount Morris, Greene Co.

Lake Erie-drainage:

Conneaut Creek, Springboro, Crawford Co.; West Springfield, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Genesee River, Rochester and Chili, Monroe Co., New York (R. H. Santens).

Black Creek, Cliili, Monroe Co., New York (R. H. Santens).

Conestogo River, Conestogo, Waterloo Co., Ontario, Canada (Miss Maria Jentsch).

Creek at North Fairfield, Huron Co., Ohio (0. E. Jennings).

Sandusky River, Upper Sandusky, Wyandot Co., Ohio (C. Goodrich).

Cedar Creek, Jerusalem Township, Lucas Co., Ohio (C. Goodrich).

Swan Creek, Toledo, Lucas Co., Ohio (C. Goodrich).

Maumee River, Roche de Boeuf Rapids, Lucas Co., Ohio (C. Goodrich).

St. Marys River, Rockford, Mercer Co., Ohio (C. Goodrich).

Blanchard River, Findlay, Hancock Co., Ohio (C. Goodrich).

Silver Creek and Beaver Creek, Williams Co., Ohio (C. Goodrich).

Ten Mile Creek, Toledo, Lucas Co., Ohio (C. Goodrich).

Otter Creek, La Salle Township, IMonroe Co., Michigan (C. Goodrich).

Raisin River, Adrian, Lenawee Co.; Grape P. 0., Monroe Co., Michigan (C. Goodrich).

Long Lake, Fenton, Genesee Co., IMichigan (C. Goodrich).

Ohio- and Mississippi-drainage:

Chautauqua Lake, Chautauqua Co., New York.^®^

Mahoning River, Leavittsburgh, Trumbull Co., Ohio (Smith collection).

I have numerous specimens collected by R. Foerster, D. Brown, D. R. Sumstine, P. E. Nordgren,
and Miss B. Ortmann, from various localities in the lake. They represent a somewhat dwarfed form,
which in this respect resembles the var. rosacea, but agrees in other respects with typical luteola.

OETMANN: monograph op the naiades of PENNSYLVANIA.

287

West Branch Nimishillen Creek, Canton, Stark Co., Ohio.

Tuscarawas River, Ohio (Holland collection).

Wolfe Creek, Wolfe Creek P. O., Washington Co., Ohio (W. F. Graham).

Scioto River, Kenton, Hardin Co., Oliio (C. Goodrich).

Ohio Canal, Columbus, Franklin Co., Ohio (Smith collection).

Ohio River, Toronto, Jefferson Co., Ohio.

West Fork River, Lynch Mines, Harrison Co.; Lightburn and Weston, Lewis Co., West Virginia.

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.

Mud River, Milton, Cabell Co., West Virginia.

Licking River, Farmer, Rowan Co., Kentucky.^®®

Muscatine Slough, Muscatine, Muscatine Co., Iowa (Hartman collection).

Mississippi River, Moline, Rock Island Co., Illinois (P. E. Nordgren).

Kishwaukee River, Rockford, Winnebago Co., Illinois (P. E. Nordgren).

West of Mississippi:

Kansas River, Lawrence, Douglas Co., Kansas (R. L. Moodie).

Bull Creek, Miami Co., Kansas (C. Goodrich, donor) (Osage-drainage).

Spring River, WiUiford, Sharp Co., Arkansas (H. E. Wheeler).

Ouacliita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler) .1®^

Fourteen Mile Creek, McBrides Switch, Wagoner Co., Oklahoma (F. B. Isely).^®^

Distrihution and Ecology in Pennsylvania (See fig. 30): This is a common
species in western Pennsylvania, although not found everywhere. It is averse
to large rivers, and, if at all present in them, is either rare, or is found under special
conditions, in smaller branches of the main river (as for instance at Neville Island).
On the other hand, it is very abundant in some of the smaller streams, as for in-
stance, in the whole Beaver-drainage. It avoids certain streams, and this is espe-
cially true of mountain-streams, such as Cheat River and the Kiskiminetas-
drainage.^®® It occurs in the only tributary of Lake Erie, in our state, which
contains shells (Conneaut Creek).

I have seen this species in Oil Creek, Orlando, Lewis Co., West Virginia.

I have also seen specimens in Fleming Creek, Pleasant Valley, Nicholas Co,, Kentucky.

Some specimens from Ouachita River and from Oklahoma are quite typical, but Wheeler (1918,
p. 117) does not report L. luteola from Clark Co., Arkansas. In this region peculiar forms turn up,
some of which develop reddish color in the nacre, and others intergrade in the direction of L. hydiana
(Lea). I omit these since I have not enough material to clear up their affinities.

The fauna of this system is poorly known. However, remarkably enough, this species has not
been listed by Harn (1891), whose material seems to have come chiefly from the Kiskiminetas-Conemaugh-
drainage. This serves to substantiate the above conclusion, since I never found this species in the two
creeks of this region, where I have been able to collect the fauna (upper Loyalhanna and Quemahoning).
Also in Yellow and Two Lick Creeks, where a fragmentary fauna has been observed, this species was
not present.

288

MEMOIRS OF THE CARNEGIE MUSEUM.

Ecologically this species is found under a number of conditions, but there is
no doubt that it prefers rather quiet water and sandy-muddy bottoms. Strong
currents and rough bottoms do not suit it, and although occasionally found in
riffles, it probably has in such cases been washed out of the quieter pools. In the
quiet water below riffles, where there is more or less muddy bottom, or in slowly
running water with fine gravel, sand, and mud, it is abundant. This accounts
for its absence in mountain-streams, and its absence or scarcity in the large rivers,
while its most favorable localities are in the plateau-streams, or the streams within
the glacial drift, which are more sluggish and have finer bottom material. For
this reason, this species is also much inclined to go into lakes.

General distribution: Type locality, According to Lamarck this species comes
from the Susquehanna and INIohawk Rivers. Its presence in the Susquehanna
has never been confirmed, but in the Mohawk it has been found subsequently
(Lewis, 1860; Marshall, 1895), and also at other places in the Mohawk and Hudson
drainages, as well as in the Erie Canal (DeKay, 1843; Simpson, 1891; Marshall).
The Mohawk should be designated as the type-locality. This is in the Atlantic-
drainage, but the chief distribution of this species is in the Interior basin.

In New York it is also knowm from the St. Lawrence-drainage (Marshall,
1895), and material in the Carnegie Museum from the Genesee system shows that
it is the typical luteola, which is found there. It is also found in New York in
the Ohio-drainage, in Ischua Creek, Cattaraugus Co., and Lake Chautauqua
(Marshall) . How far the typical form extends northward in this region is unlmown.
It has been reported from Canada (Ontario, and also westward in the St. Lawrence-
drainage up to Lake Superior, and from the Hudson Bay-drainage) but in these
northern parts it seems, that its place is largely, if not entirely, taken by certain
varieties, chiefly rosacea (See below). Yet the typical form is certainly present
north of Lake Erie as is shown by specimens in the Carnegie Museum (Conestogo
River) (also reported from northern Michigan, in Manistique River, Schoolcraft
Co., by Winslow, 1917).

Westwards L. luteola is found practically aU over western Pennsylvania,
Ohio, Indiana, and Illinois, crossing over into the lake-drainage at various points,
and it extends northward through Iowa, Wisconsin, and Minnesota into the
drainage of the Red River of the North, although, as has been said, it seems in
the northern parts of its range to give way to varieties.

Lewis (1877) calls attention to the fact that this species has not been recorded
from “any stream much south of the latitude of the Ohio River,” but that possibly
it is present in Kentucky. Material in the Carnegie Museum shows that it occurs

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289

in West Virginia not only in the headwaters of the Monongahela, but also in the
Little Kanawha, in the Kanawha and Guyandotte drainages ; but that it is absent
here in certain regions, for instance, in the Elk River of the Kanawha system.
This no doubt is due to the rough character of Elk River. In addition I found
this species in the Licking River in Kentucky. But it actually seems to be absent
in the Cumberland and Tennessee River drainages.

In a westward and southwestward direction this species undoubtedly goes as
far as Missouri (Utterback, 1916), Kansas (Scammon, 1906), Arkansas, and Okla-
homa, and has been reported to exist in Texas (Brazos River, Simpson, 1900).
However, in this region it passes into other forms {hydiana, for instance) which
may, or may not be, “good species.” In Mississippi and Alabama it seems to be
represented by kindred forms, which are generally considered to be different
species.

Thus L. luteola has its main range chiefly in the northern section of the Interior
Basin, north of the Ohio, and passes into the northern drainages of the St. Lawrence
and Hudson Bay. But in the north, as well as in the south, it inclines to variation,
and has developed geographical and local races, some of which may be regarded
as valid species.

Both Baker (1898a) and Scammon (1906) call this a mud-loving species, which
agrees with my observations. Although sometimes found in other environment,
it thrives best and is most abundant in mud and sluggish or quiet water. This
satisfactorily explains its absence in certain streams in western Pennsylvania and
West Virginia, and possibly also in the Tennessee-Cumberland-drainage. This
also holds good in the Ohio River between Pennsylvania and Cincinnati. I found
it in the Ohio at Toronto, in a small branch of the river, but at no other locality,
and it was entirely missing in the numerous piles of the clam-diggers examined by
myself.

Lampsilis luteola rosacea (DeKay) (1843).

Lampsilis luteola rosacea (DeKay) Simpson, 1914, p. 62; Walker, 1913, p. 21.

Plate XVH, figs. 3, 4, 5.

Record from Pennsylvania:

Ortmann, 1919&, p. 202 (as lake-form of L. luteola).

Characters of variety: This may be described as a small, stunted form of L.
luteola, generally with rather light-colored (pale yellow to rusty) epidermis, with
white or (locally) rosaceous nacre. The specimens from Lake Erie (and also other
lakes) have generally more regular, more crowded, and more distinct growth-rests.

290 MEMOIRS OF THE CARNEGIE MUSEUM.

L. H. D.

Size: (Males) 1. Erie, Cat. No. 61.4858 89 trim. 49 mm. 31 mm.

2. do. Cat. No. 61.4020 76 “ 44 “ 27 “

3. do. Cat. No. 61.4020 51 “ 29 “ 18 “

(Females) 4. Erie, Cat. No. 61.4020 (gravid) 81 “ 54 “ 35 “

5. do. Cat. No. 61.4020 (gravid) 64 “ 41 “ 28 “

6. do. Cat. No. 61.4860 52 “ 34 “ 20 “

Soft parts identical with those of the main species, glochidia also identical.

Breeding season: Gravid females are on record for May 21, 1909; May 22,
1909; May 24, 1909; July 7, 1910; July 12, 1910. All these specimens had
glochidia, and on July 7 and 12 a number were observed discharging the glochidia.
The variety is certainly hradytictic, like the main species.

Remarks: The original description of rosaceus DeKay is founded upon speci-
mens with special features: rosy nacre and pallid brown epidermis. But Simpson
(1900, p. 535, footnote 1) pointed out that this probably is only a local effect,
and that other specimens have white nacre and differently colored (darker) epi-
dermis.^^® I find, that there is in Lake Erie great uniformity in the color of the
nacre, which is always white, while there is great variability in the color of the
epidermis. Specimens from the surf-beaten shores of the open lake are remarkably
light, and the ground color of the epidermis is light yellow or light greenish (grayish-
green), while those from protected localities in Presque Isle Bay, and from beach-
pools, are more greenish-olive, lighter or darker, and often incline toward a reddish
brown, which is most intense and brilliant towards the beaks. In the gTowth-
lines there is great variability, but the tendency is to have them rather closely set,
rather distinct and regular. This is most evident in specimens from deeper water,
least so in those from the beach-pools.

The characters of this variety are very poorly marked. It is true that speci-
mens from Lake Erie generally are recognizable by size, color, and distinct growth-
lines; but in specimens from other localities these characters are more or less
unstable, and transitional conditions toward true luteola are frequent. I have
already mentioned that in L. luteola from Lake Chautauqua, which incline toward
rosacea in size, other characters are normal. Specimens from Winona Lake look
much like the Lake Erie form, having the reddish epidermis and very distinct
growth-lines, but are larger. Specimens from Moose River are small and pale in
color, but the growth-lines are indistinct.

Specimens collected by O. E. Jennings on the North shore of Lake Superior
much resemble the Lake Erie form, being pale in color, without rays, but they differ

Utterback (1916, p. 185) relying only on the color of the nacre, erroneously called the red-nacred
form of Missouri (also found in Arkansas) by the name of L. luteola rosaceus.

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291

in different localities. A specimen from Nipigon Straits is absolutely like the
Lake Erie form, while others from Six Mile Lake (a shallow lake on Thimdercape
Peninsula) have irregular growth-lines, and incline more toward normal luteola.
These specimens could not very well be called superior ensis, for they do not have
a dark epidermis. Then again specimens from Kaministiquia River (tributary
to Lake Superior at Fort William) are darker in color, and might possibly represent
the real superior ensis.

Specimens from Sheyenne River, North Dakota, are indistinguishable from
certain Lake Erie specimens, except for the irregular growth-lines.

I have the impression that rosacea is not so much a geographical, as an eco-
logical race, produced by the environment of great lakes, and that it turns up,
whenever the proper conditions are offered. Since this environment is very widely
distributed in the northern states and in Canada, rosacea appears as a geographical
race. In this connection it should be emphasized that this variety is not found
in the tributaries of the lakes, but that the normal form of luteola exists there.
There are also certain lakes, in which the form rosacea is not present, for instance
Lake Chautauqua, and Conneaut Lake. In the latter luteola is quite typical.

More detailed investigations are required to finally determine this question.
It also should be ascertained to what factors the rose-colored nacre may be due.
Our specimens from Sandy Lake, Ontario, have red nacre, and come from a lake,
the water of which is, according to Mr. G. H. Clapp, heavily charged with lime.

Localities in Pennsylvania represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, outer shore, and beach-pools of Presque Isle, Erie, Erie Co.

Other localities represented in the Carnegie Museum:

Lake Ontario, Braddock Bay, Manitou Beach, Monroe Co., New York (R. H. Santens).

Lake Erie, Sandusky Bay, Cedar Point, Erie Co., Ohio (C. Brookover) (0. E. Jennings); La Plaisance
Bay, Monroe Co., Michigan (C. Goodrich); Crystal Beach (F. Behrle) and Port Colborn, Welland
Co. (C. Goodrich); Port Rowan and Port Dover, Norfolk Co., Ontario, Canada (C. Goodrich).
Sandy Lake, Peterboro Co., Ontario, Canada (G. H. Clapp).

Grand River, Cayuga, Haldimand Co., Ontario, Canada (C. Goodrich).^'*''

Winona Lake, Kosciusko Co., Indiana (E. B. Williamson) (not typical).

Lake Wiwassee, Kosciusko Co., Indiana (C. Goodrich).

Lake Huron, Port Huron, St. Clair Co., Michigan (N. M. Grier); Saginaw Bay, Michigan (Smith col-
lection).

Big and Little Whitefish Lakes, Pierson, Montcalm Co., Michigan (Miss M. O’Malley).

Au Sauble Lakes, Lake Co., Michigan (C. Goodrich).

Douglas Lake, Cheboygan Co., Michigan (H. B. Baker).

A small race of luteola, and probably var. rosacea, although not typical. It should be noted
that in the Upper Grand River-drainage (Conestogo River), the real L. luteola is present.

292

MEMOIRS OF THE CARNEGIE MUSEUM.

Crystal Lake, Benzie Co., Michigan (B. Walker, donor).

Lake Michigan, off Kenosha, Kenosha Co., Wisconsin (P. E. Nordgren).

Lake Superior, Nipigon Straits, St. Ignace Island, Ontario, Canada (0. E. Jennings).

Six Mile Lake, Silver Island, Thundercape Peninsula, Ontario, Canada (0. E. Jennings).

Lake Nipigon, Ombabika Bay, Ontario, Canada (0. E. Jennings).

Sheyenne River, Argusville, Cass Co., North Dakota (S. M. Edwards).

Moose River, South of Moose Factory, Ontario, Canada (W. E. C. Todd).

Distribution and Ecology (See fig. 30) : Type locality, Seneca Lake, New York
(DeKay).

As Simpson says, this race is found in the St. Lawrence area, and is common in
the lakes. However, its exact distribution is not very well known, since it often has
not been kept apart from the typical form. In addition our records show, that, in
northern Indiana, it is also in some lakes belonging to the Ohio (Wabash) drainage.

Walker (1913) has it from Lake Erie, and he was the first to use the name
rosacea for the lake-form. Sterki (1907a) reports it from Lake Erie in Ohio, but
he also gives typical luteola from this lake. It may be, that he understood by
rosacea only specimens with red nacre. But I have never seen such from Lake
Erie. Walker (1898) reports borealis and superiorensis from northern Michigan,
but not rosacea. What I have from Lake Huron and central and northern Michi-
gan, is indistinguishable from the Lake Erie form. This variety certainly also
occurs in Lake Michigan and Lake Superior.

A form very close to this is found in Aloose River, near Hudson Bay. It also
turns up again, although not quite typical, in the drainage of the Red River of the
North in North Dakota.

The ecological preference of this form is certainly for the lake environment,
and it is best developed in the larger lakes. In Presque Isle Bay in Lake Erie,
where it is a common shell, it is found everywhere on the sandy and gravelly shores,
in shallow water, and down to a depth of about fifteen feet in sand and mud. It
also is found in the beach-pools of Presque Isle, upon sandy-muddy bottom, and
is one of the few shells existing in the open lake, being frequently thrown out alive
by the surf.

Lampsilis radiata (Gmelin) (1792).

Lampsilis radiata (Gmelin) Simpson, 1914, p. 64.^®^

Plate XVII, figs. 6, 7.

Records from Pennsylvania:

Gabb, 1861 (Schuylkill and Wissahickon, and League Island, Philadelphia).

Bruckhart, 1869 (Lancaster Co.)

The figures given by DeKay (1843, PI. 19, figs. 237, 238) as of Unio ochraceus, have always been
taken for L, ochracea (Say), even by Simpson (1914, p. 50), but they actually represent L. radiata, and

ORTMANN: monograph of the naiades of PENNSYLVANIA.

293

Hartman & Micliener, 1874 (Schuylkill River, Chester Co.).

Dean, 1891 (West Branch Susquehanna River, Muncy, Lycoming Co.).

Pilsbry, 1894 (Susquehanna River, York Furnace, York Co.).

Schick, 1895 (Delaware River, Philadelphia; Canal at Manayuiik).

Ortmann, 19095, p. 204.

Characters of the shell: Shell of about the same size and similar in shape to
that of L. luteola. Beak-sculpture also similar (Marshall, 1890, fig. 4), but valves
much more compressed. Epidermis yellowish, or light or dark green, more rarely
inclining to brownish, not smooth, but roughened by close concentric wrinkles.
Dark green or blackish rays are generally present, but they are not well-defined,
except that in light-colored specimens they are sharper. Concentric bands of
lighter and darker color are often present.

Hinge similar to that of L. luteola, as well as the rest of the inside of the shell,
but the color of the nacre is more variable, sometimes being entirely white, often
tinted with pinkish or salmon, and even entirely of these colors.

Sexual differences of the shell much less marked than in L. luteola. The female
shell has a similar tendency to expand in the postbasal region, but only slightly,
and never so distinctly as in L. luteola. Some females are even hard to distinguish

from males.

L. H. D.

Size: 1. Manayunk, Cat. No. 61.4051 (probably cf) 93 mm. 53 mm. 27 mm.

2. do. Cat. No. 61.1789 (9 ) 88 “ 52 “ 28 “

3. Tunkhannock, Cat. No. 61.4054 (gravid 9) 87 “ 49 “ 29 “

4. Yardley, Cat. No. 61.3473 (probably cf ) 71 “ 42 “ 21 “

5. Selinsgrove, Cat. No. 61.4868 (d") 70 “ 40 “ 23 “

6. South Waverly, Cat. No. 61.4053 (9) 60 “ 34 “ 17 “

Soft parts (See Ortmann, 1912, p. 349) figured by Lea (Obs. II, 1838, PL 15,
figs. 48, 49). Glochidia (See Lea, Obs. VI, 1858, PL 5, fig. 20). I observed them
in a specimen from Severn River, Ontario. They agree with those of L. luteola:
L. 0.22 to 0.23, H. 0.27 to 0.28 mm.

Breeding season: According to Conner (1907) “all the year round.” I have
taken a gravid female with eggs on Aug. 22, 1909. This would indicate the begin-
ning of the season. The female with glochidia was collected Aug. 20, 1914. It
might be that this represents the end of the breeding season, which thus would
overlap with the next one.

are both probably females, which is seen at once by the general outline of the shell. It is remarkable,
that this mistake of DeKay has never been discovered, although his error with reference to his Unio
radiatus (which is Eurynia iris novi-eboraci) has been corrected previously. There is no doubt that the
confusion within the two species, radiata and ochracea, is in large part due to these incorrectly named
figures of DeKay.

294

MEMOIRS OF THE CARNEGIE MUSEUM.

Remarks: The distinctness of this species from L. luteola has been much
discussed. Some writers (Dean, 1891) hold that it is sharply separated, while
others, among them Simpson (1891), admit that there apparently are intergrades.

According to my experience I must say, that I never had in our state any
trouble in distinguishing the two species. L. radiata is more compressed, lacks
the shining epidermis, often has reddish nacre, and the difference between the male
and the female shell is considerably less marked than in L. luteola.

But I possess a few specimens which indeed are somewhat abnormal. This
applies chiefly to some I received from C. H. Conner, and which come from Newton
Lake, Camden Co., New Jersey. These are more swollen than the normal form,
and in the female the shell is more expanded in the postbasal region, and in shape
they thus greatly resemble L. luteola. However, even in these specimens the epi-
dermis has the fine wrinkles of L. radiata, and is not smooth and shining as it is
in L. luteola. I think that these specimens represent the reaction of L. radiata to
the lake-environment.

With respect to shape and principally color, this species appears to be quite
variable outside of Pennsylvania, but within this state it is rather uniform. It
seems to me that in the northern parts of its range lighter epidermis (yellow to
light brown) prevails, while farther south darker tints (dark green to brownish)
are the rule. In the extreme south (southern Virginia and North Carolina) this
species seems to develop a distinct local race (var. conspicua (Lea)).

It should be mentioned, that L. radiata to a certain degree resembles Acti-
nonaias ligamentina in general shape and color, and the dark green specimens
with broad rays distinctly recall the latter species. Such specimens may be recog-
nized by the more elongated shape, thinner shell, and by the rough epidermis.
Likewise the presence of pink in the nacre, and the beak-sculpture serve to dis-
tinguish them.^^^

Of course, the examination of the soft parts always shows that these two species
have no close genetic relationship.

Localities in Pennsylvania represented in the Carnegie Museum:

Delaware River, Yardley, Bucks Co.

Schuylkill Canal, Manajuink, Philadelphia Co.

Susquehanna River, Selinsgrove, Snyder Co.

West Branch Susquehanna River, Williamsport, Lycoming Co. (D. A. Atkinson).

North Branch Susquehanna River, Tunkhannock, Wyoming Co.

A specimen from Grand Rapids, Michigan, in the Carnegie Museum, was labeled "U. radiatus,
and, has indeed, much external resemblance to this. But closer inspection has shown that it is Actinonaias
ligamentina, having the smooth epidermis and the beak-sculpture of this species.

ORTMANN; monograph of the naiades of PENNSYLVANIA.

295

Chemung River, South Waverly, Bradford Co.

Other localities represented in the Carnegie Museum:

Ottawa, Ontario, Canada (Hartman collection) (marked: “ Lea datum ”)•

St. Lawrence River, Bluff Island, Clayton, Jefferson Co., New York (Miss A. H. Robinson) (H. Kahl);

Grindstone Island, Clayton, .lefferson Co., New York (H. Kahl).^®^

Severn River, Gloucester Pool, Muskoka Co., Ontario, Canada (0. A. Peterson).

Spider Bay of Georgian Bay, Sans Souci, Parry Sound, Ontario, Canada (H. Kahl).

Connecticut River (Hartman collection).

Herldmer Co., New York (Smith collection).

Keuka Lake, Yates Co., New York (G. H. Clapp, donor) (Lake Ontario-drainage).

Little Lakes, Herkimer Co., New York (Smith collection).!®^

Newton Lake, Camden Co., New Jersey (C. H. Conner) (See above).

Delaware River, Newbold, Gloucester Co. (C. H. Conner); and Fish House, Camden Co., New Jersey.
Potomac River,- Washington, D. C. (G. H. Clapp, donor).

Distribution and Ecology in Pennsylvania (See fig. 31) : In Pennsylvania this
species is found in the Delaware and Susquehanna drainages, but it is absent in

• Lampsilis radiata.
m Lampsilis cariosa.
+ Lampsilis ochracea.

the smaller streams which flow to the Potomac. Altogether it is rare according
to my experience, but this may be due to the fact that I did not do much collecting

A dwarf form, with light brown ground-color of epidermis. These specimens might represent
U. borealis Gray.

This is in Herkimer Co., not in Otsego Co., as given by Lewis (1856, p. 259). These lakes drain
into Otsego Lake, and this in turn into the Susquehanna River.

296

MEMOIRS OF THE CARNEGIE MUSEUM.

ill the tidewaters of the Delaware. According to the older records, its metropolis
seems to be in the lower Delaware near Philadelphia. It ascends the larger rivers
(Delaware and Schuylkill), but the upper boundary has not been ascertained.

In the Susquehanna-drainage it is distinctly a rare shell. It has been found
in the lower part (York Haven, Pilsbry), but otherwise it is restricted to its larger
branches. In the West Branch it has been traced up to Williamsport, in the
North Branch to the New York state line.^^®

Although present in the lower Potomac, it has never been found in the drainage
of this river in Pennsylvania, and it seems to be absent in the whole upper Potomac
system West of the Blue Ridge Mountain.

Having found this sjiecies only at a few places, and probably not in the most
favorable environment, I am unable to say what are the ecological conditions,
which it prefers. In the Delaware above Trenton and in the Susquehanna I found
it in strongly flowing water in gravel. But here it was unquestionably rare.
If its metropolis is in the dower parts of the large rivers, it might be a tidewater
species, prefering quiet water, and sandy or muddy bottom. At any rate, I found
it on sandy bottom in the lake-like part of the Delaware at Fish House, New
Jersey.

General distribution: Type locality, Given by Gmelin from Malabar, which,
of course, is incorrect. Lamarck (1819) reports it from Saratoga Lake in New
York, and if there should not be any other earlier record, we might select this as
the type locality. Simpson (1914) gives Virginia as type locality.

This species belongs to the Atlantic-drainage from Virginia to Maine. In
Virginia it is chiefly present according to Conrad (1836) in the tidewaters,^®® and
farther South it assumes a different shape, and has been called conspicua. Its
presence in the lower Potomac in Virginia and the District of Columbia is well
established (Dewey, 1856 ; Alarshall, 1895) . It is known from the state of Delaware
(Rhoads, 1904), Pennsylvania, New Jersey (lower Delaware River, and Second
River, Belleville, Essex Co., as ochraceus De Kay, 1843), from New York, Con-
necticut, Rhode Island, JMassachusetts and IMaine (See Linsley, 1845; Perkins,
1869; Gould-Binney, 1870; Carpenter, 1890; Lermond, 1909; Johnson, 1915).
In New York (Marshall, 1895) it ascends the Atlantic rivers (Upper Susquehanna
and Hudson) reaching the Mohawk, and crossing over into the St. Lawrence-
drainage, where it is found in Lake Ontario and its tributaries, and down the

It goes farther up in New York state, since it occurs in Little Lakes and Schuyler’s Lake in the
region of the headwaters, and has been reported from Chenango and Tioga Rivers.

It is positively absent in the mountains west of the Blue Ridge, and also seems to be absent or
rare on the Piedmont Plateau.

OKTMANN: monograph of the naiades of PENNSYLVANIA. 297

St. Lawrence to Ottawa and Montreal (Bell, 1859; Simpson, 1891; Marshall,
1895).

Possibly it extends its range westward from the lower St. Lawrence, for it
has been reported from Lake Nipissing (Bell, 1859), and even farther West, from
Lake Superior, Lake Winnipeg, and Nelson River, but these latter localities require
confirmation, and a few additional records from Ohio and Indiana are certainly
wrong. Its presence in Michigan is doubtful (Walker, 1898). But it is positively
present in the region of Georgian Bay: specimens in the Carnegie Museum are
nearly normal, except that the color is often more brownish.

This is an Atlantic species, belonging to the northern element in the Atlantic
fauna (Ortmann, 1913a). In New York it has crossed over into the lower St.
Lawrence-drainage, and apparently has here a tendency to spread westward
along a route, which lies North of Lake Erie. If this should prove to be correct,
attention should be called to the similarity of this range with that of Elliptio
violaceus.

In New York its range in part overlaps that of L. luteola (certainly in the
Mohawk and St. Lawrence drainages), but particulars as to the mutual relation
and possible association of these two species are lacking. This is a question which
requires closer investigation.

As far as known, this species prefers tidewaters, but ascends some of the
larger rivers to a considerable distance (Hudson, Susquehanna). Nevertheless in
the south it apparently does not have this tendency. The fact that it often goes
into canals and is frequently found in lakes, also indicates that rough water and
rough bottom (riffles) are not very favorable to it.

Lampsilis ovata (Say) (1817).^^^

Lampsilis ovatus Simpson, 1914, p. 48.

Plate XVII, figs. 8, 9; Plate XVIII, figs. 1, 2, 3.

Records from Pennsylvania:

Call, 1885 (Allegheny River, up to central New York).

Clapp, 1895 (Allegheny Co.).

Marshall, 1895 (Allegheny River, Warren, Warren Co.).

Rhoads, 1899 (confused with L. ovata ventricosa, but specimens from Ohio River, Coraopolis, Allegheny
Co., and Beaver, Beaver Co., belong here).

Ortmann, 19096, p. 189.

Characters of the shell: For reasons given below, it is best to describe this
form in terms of comparison with its variety, L. ovata ventricosa (See remarks).

1” Not 1816.

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MEMOIRS OF THE CARNEGIE MUSEUM.

Similar to L. ovata ventricosa, but distinguished by the development of the
jiosterior ridge, which is very sharp and very distinct toward the beaks. In
consequence of this the posterior slope is flattened, and in most cases even concave,
chiefly towards the beaks. Further towards the anterior end of the shell the two
valves are not uniformly convex, but peculiarly compressed, rendering the hori-
zontal cross-section of this part wedge-shaped.

In addition there is a tendency toward the suppression of the rays. Rays are
indeed sometimes present, but in most cases the epidermis is uniformly greenish
yellow, dark upon the posterior slope and without rays. Old males become drawn
out at the posterior end, almost rostrate.

L. H. D.

Size: (Males) 1. Aladdin, Cat. No. 61.3426 136 mm. 92 mm. 57 mm.

2. Walnut Bend, Cat. No. 61.3423 103 “ 72 “ 46 “

3. Godfrey, Cat. No. 61.4009 86 “ 61 “ 37 “

(Females) 4. Industry, Cat. No. 61.3425 126 “ 94 “ 64 “

5. Cambridge Springs, Cat. No. 61.4010 (gravid) .100 “ 81 “ 48 “

6. Cochranton, Cat. No. 61.3420 83 “ 64 “ 39 “

Soft parts (See Ortmann, 1912, p. 350, fig. 26). Glochidia (See Lea, Obs. VI,
1858, PL 5, fig. 15). I have found them to measure: 0.24 X 0.28 mm.

Breeding season: Aly records for gravid females cover the period from August
4 to October 24, and from May 23 to May 25. The spring records are scanty, but
the species is surely hradytictic, and there is an interim at least in July.

Remarks: There are numerous intergrades between L. ovata and L. ventricosa.
In the larger rivers, the two forms are practically always associated, and the
transition from a very sharp to an almost entirely effaced posterior ridge with
the corresponding intermediate condition of the posterior slope, is frequently found.
Likewise the other characters given above pass into each other. In color there is
also great variability, and, although the true ovata has generally no rays, rays are
sometimes very well-developed; on the other hand in true ventricosa the rays may
be lacking. I have quite a number of specimens, which are so completely inter-
mediate, that I am unable to assign them to either form.

Going up beyond a certain point in our rivers, and into the smaller tributaries,
L. ovata disappears, and its place is entirely taken by L. ovata ventricosa, and there
are many creeks, where only the form ventricosa exists with no trace of ovata, and
no indications of an inclination toward it.

Exactly the same conditions prevail in the upper Tennessee-drainage in
eastern Tennessee and southwestern Virginia, and a form corresponding to L.
ovata ventricosa develops out of the true L. ovata in the headwaters. However, the

ORTMANN: monograph of the naiades op PENNSYLVANIA.

299

Tennessee ventricosa has some peculiarities of its own. These shells will be treated
elsewhere.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Industry, Beaver Co.; Coraopolis (S. N. Rhoads), Neville Island, and Edgeworth, Allegheny
Co. (G. H. Clapp).

Allegheny River, Braeburn, Westmoreland Co.; Aladdin, Godfrey, Johnetta, Kelly, and Templeton, Arm-
strong Co.; Walnut Bend, Venango Co.; Tionesta and Hickory, Forest Co.; Warren, Warren Co.
French Creek, Cochranton, Meadville, and Cambridge Springs, Crawford Co.

Conneaut Outlet, Conneautlake, Crawford Co.

Other localities represented in the Carnegie Museum:

Ohio-drainage:

Ohio River, Toronto, Jefferson Co., Ohio; Wheeling, Ohio Co., West Virginia (W. F. Graham); St.

Marys, Pleasants Co., West Virginia; Portland, Meigs Co., Ohio.

Elk River, Shelton and Clay, Clay Co., West Virginia.

Tennessee-drainage :

Tennessee River, Tuscumbia, Colbert Co., and Florence, Lauderdale Co., Alabama (H. H. Smith).

Paint Rock River, Paint Rock and Princeton, Jackson Co., Alabama (H. H. Smith).

Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Alill and Holston
Station, Grainger 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.

Distribution and Ecology in Pennsylvania (See fig. 32) : This species is abundant
in the Ohio below Pittsburgh, and in the Allegheny all the way up to Warren
County. According to Call (1885) it passes by this route into New York, but
Marshall (1895) does not report it from the uppermost Allegheny and it has not
been found in McKean County, Pennsylvania. From the Allegheny it enters
French Creek and extends up to Cambridge Springs. It also enters Conneaut
Outlet (but only one dead shell was found there). It is found in no other stream,
and its absence is especially noticeable from the whole Beaver-drainage, where
L. ovata ventricosa on the other hand is common.

No records from the Monongahela are known. However, it is probable that
it once existed in the Monongahela proper, at least as far as to the West Virginia
state line, for I was able to identify it among the shells from an Indian garbage
heap at Point Marion (Ortmann, 1909c). It is not found in the upper Monon-
gahela-drainage in West Virginia, where L. ovata ventricosa is locally abundant.
Thus L. ovata is clearly a form prefering larger rivers, and, according to my

300

MEMOIRS OF THE CARNEGIE MUSEUM.

observations, it inhabits in these the roughest parts, riffles with strong currents,
the bottoms consisting of large stones, loosely piled over each other, with little
finer material packing them together. Of course, occasionally it is found also in
finer gravel. I have no doubt, that its chief characters are reactions to the con-
ditions iirevailing in riffles. The flattened or concave, truncate posterior slope is

• Lampsilis ovata. O Indian garbage heap.

■ Lampsilis ovata ventricosa.

+ Lampsilis ovata canadensis.

produced by the current and the material rolled by it over the posterior end of the
shell, when it is imbedded in the gravel; and the wedge-like attenuation of the
anterior end is a device to enable the shell to plough through the heavy gravel in
which it lives. I have seen the shell moving along in coarse gravel, and pushing
aside stones, which were much larger than itself.

General distribution: Type locality^ Ohio (Say).

It seems that the Ohio River, the Cumberland, and the Tennessee represent
the metropolis of this species. Aside from western Pennsylvania, it is found in
the Ohio proper in West Virginia (Carnegie Museum), Ohio (Sterki, 1907a),
Indiana. (Call, 1896a and 1900), and Illinois (Baker, 1906). It occurs in very few
of the tributaries. Hildreth (1828) gives it from the Muskingum River at Marietta,
Washington Co., Ohio; Sterki (1907a) from the Great Miami River; and Call
from the Wabash, White, and Eel Rivers. In addition, I have found it in Elk
River in West Virginia, but only as far up as Clay County. Above this point,
its place is taken by ventricosa, and the mutual relations of these two forms are

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301

exactly as elsewhere. In the Cumberland, it goes up to the falls (and a little
beyond), according to Wilson & Clark (1914); and it has been reported from the
Tennessee River as far up as the Holston (Lewis, 1871; Call, 1885), and here it
reaches southwestern Virginia, as I have ascertained, in the Clinch River.

The distribution is thus rather restricted, being confined to the larger rivers
of the Ohio system. Outside of Pennsylvania I found this species in the rough
parts of rivers. In the Ohio below Pittsburgh, where there is more steady current
and finer bottom material, it is by no means so abundant as in the Allegheny,
for instance. The clam-diggers on the Ohio take it regularly, but not in great
numbers.

I have treated as spurious a few records of this species from the lower St.
Lawrence in Canada and the Maumee River.

Lampsilis ovata ventricosa (Barnes) (1823).

Lampsilis ventricosa (Barnes) Simpson, 1914, p. 38.^®^

Plate XVIII, fig. 4; Plate XIX, figs. 1, 2, 3.

Records from Pennsylvania:

Ham, 1891 (western Pennsylvania) (as U. occidens and U. subovatus) .

Stupakoff, 1894 (Allegheny Co.) (as U. cariosus).

Marshall, 1895 (Allegheny River, Warren Co.) (as U. occidens).

Rhoads, 1899 (specimens from Ohio River, Coraopolis, Allegheny Co., and from Beaver River, Wampum,

Lawrence Co., recorded as U. ovatus, belong here).

Ortmann, 19095, p. 182, 202.

Characters of the shell: Shell large, often very large, rather thin when young,
but attaining considerable thickness when old. Outline subelliptical or subovate,
rather short and high, hardly ever one-and-a-half times as long as high, mostly
much shorter. Anterior margin rounded. Lower margin curved. Upper margin
short, nearly straight, passing in a blunt angle or gentle curve into the obliquely
descending posterior margin. Posterior and lower margins meeting in a blunt,
rounded posterior point. Beaks more or less swollen, moderately elevated, located
anterior to the middle of the shell. Beak-sculpture consisting of four or five rather
coarse bars, of which the second and third have a slight tendency to fall into two
loops, with a light sinus in the middle, while the first is indistinct, and the fourth

According to Vanatta (1915, p. 551), Lampsilis cardium Rafinesque (1820) is this. I do not
accept this identification, since the original description of Rafinesque clearly shows that L. cardium is
the female of L. ovata (Say).

The figures of Unio cariosus given by DeKay (1843, PI. 21, figs. 243, 244), quoted also by Simpson
(1914, p. 44) under L. cariosa, are undoubtedly L. ovata ventricosa, and not cariosa.

302

MEMOIRS OF THE CARNEGIE MUSEUM.

and fifth tend to become obsolete. All the bars are effaced on the posterior
slope. Valves more or less regularly convex, more flattened upon the sides. Pos-
terior ridge poorly developed, mostly quite obsolete, and indicated only by a
stronger convexity of the surface, but sometimes this ridge may become more
distinct, chiefly so toward the beaks. Posterior slope gently convex or almost
flat, but not concave.

Epidermis varying from lighter or darker yellowish green, to olive-brown;
smooth and shining. Rays rarely absent, mostly present, but extremely variable.
They are dark green to blackish, straight and continuous, finer or broader, and may
cover the whole surface or only part of it. On the posterior slope the epidermis
is less smooth, and generally darker. Concentric bands may be jiresent.

Hinge well developed. Pseudocardinals generally two in each valve, but
extremely variable in shape and size. In young specimens they are more or less
compressed; in older ones, more stumpy and ragged. Interdentum absent or
narrow. Laterals lamellar, high and strong, that of the right valve generally very
broad and suddenly truncated posteriorly. Beak-cavity moderate. Dorsal mus-
cle-scars in the beak-cavity. Adductor-scars distinct, rather well-impressed,
chiefly so the anterior ones. Nacre silvery or bluish white, often with a pink blush,
or more or less suffused with pink or purplish, but never entirely red.

Sexual differences of the shell well-marked. In the male, the lower margin
is rather regularly curved, and the shell has a blunt posterior point. In the female
the lower margin is considerably expanded in the postbasal region, the anterior
portion thus becoming almost straight, while the posterior ascends suddenly to
the blunt posterior end of the shell. The whole posterior section of the shell is
much higher than in the male, more expanded, and broadly rounded.

L. H. D.

Size: (Males) 1. New Galilee, Cat. No. 61.2143 155 mm. 120 mm. 76 mm.

2. Meadville, Cat. No. 61.3392 115 “ 74 “ 43 “

3. Cannelton, Cat. No. 61.2881 76 “ 52 “ 32 “

(Females) 4. Neville Island, Cat. No. 61.1582 123 “ 88 “ 56 “

5. New Galilee, Cat. No. 61.3236 (gravid) 97 “ 71 “ 46 “

6. Darlington, Cat. No. 61.2918 (gravid) 89 “ 64 “ 40 “

Soft parts (See Ortmann, 1912, p. 35) figured by Lea (Obs. VII, 1860, PL 30,
fig. 107; Ortmann, 19116, pp. 319, 320, figs. 7, 8). Glochidia (See Lea, Obs. VI,
1858, PI. 5, fig. 13; Ortmann, 19116, PL 89, fig. 23; Surber, 1912, PL 2, fig. 24).
My measurements are: 0.25 X 0.29 mm.; while Surber gives: 0.205 X 0.255 mm.
This is a rather unusual discrepancy.

Breeding season: I have a very complete series of dates for gravid females,

OETMANN: monograph of the naiades of PENNSYLVANIA,

303

from July 30 to October 24, and from May 9 to July 8. The form is bradytictic,
two succeeding breeding season approaching each other closely in July, but an
interim is apparently present in this nionth. However, an occasional overlapping
of the seasons is not impossible. Surber’s notes (1912, p. 7) on the presence of
giochidia are fragmentary, but they also indicate a possible overlapping of the
seasons.

Remarks: The taxonomic relation of this form to L. ovata has been discussed
above. Both are forms of the same species connected by numerous intergrades,
but locally they may be pure. The true ventricosa is easily recognized by size,
shape, and color. It differs from the allied L. fasciola in size, and in the character
of the rays (not interrupted or wavy). From the eastern representatives, L.
cariosa and L. ochracea it differs chiefly in size and color. L. orhiculata has a
much heavier shell and different color.

L. ovata ventricosa is very variable in thickness, shape, and color. There is
more or less tendency to form local races. In some creeks it attains giant pro-
portions, while in others it has only a medium size. The latter is the case in the
mountain-streams (Cheat, upper Loyalhanna, Quemahoning) .

The typical characters, most of all the color, are best developed in shells of
medium size. In the upper Ohio greenish is the prevailing tint of the epidermis,
which is also glossy; though in old shells the epidermis frequently is discolored
and loses its gloss, such shells appearing dull brownish or even blackish. On the
average the greenish epidermis and the dark rays give to the shell a comparatively
dark aspect, but specimens with light yellowish gveen or light brownish epidermis,
and without rays, are not infrequent. If the nacre is strongly tinted with pink,
this color sometimes influences the outer color in young specimens, which appear,
reddish brown. A peculiar local race has developed in Lake Erie, and will be
discussed below.

With regard to the convexity of the valves and the inflation of the beaks, the
Pennsylvanian form varies a little, but not so much as in the southern and western
sections of its range. In the South, it is represented by a form with much inflated
beaks {satura Lea) which also often has a very dark blackish epidermis.

Localities in Pennsylvania represented in the Carnegie Museum:

The Ohio and its smaller tributaries:

Ohio River, Smith’s Ferry, Cook’s Ferry, Shippingport, and Industry, Beaver Co.; Coraopolis (S. N.

Rhoads), Neville Island, and Edgeworth (G. H. Clapp), Allegheny Co.

Little Beaver Creek, Cannelton (Miss Vera White; H. H. Smith), Darlington, and New Galilee, Beaver

Co.; Enon Valley, Lawrence Co.

304

MEMOIRS OF THE CARNEGIE MUSEUM.

Raccoon Creek, New Sheffield, Beaver Co.

Chartiers Creek, Carnegie, Allegheny Co. (D. A. Atkinson).

Beaver-drainage:

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Ellwood City, Lawrence Co. (G. U. Clapp & H. H. Smith); Harmony, But-
ler Co.

Slipperyrock Creek, Wurtemberg and Rose Point, Lawrence Co.

Mahoning River, IMahoningtown and Hillsville, Lawrence Co.

Neshannock Creek, Eastbrook, and Volant, Lawrence Co.; Leesburg, Mercer Co.

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.; Clarksville, Shenango, and Jamestown,
Mercer Co.

Pymatuning Creek, Pymatuning Township, IMercer Co.

Little Shenango River, Greenville, Mercer Co.

Alleghemj-drainage:

Allegheny River, Natrona, Allegheny Co.; Aladdin, Godfrey, Johnetta, Kelly, Mosgrove, Templeton,
and Parkers Landing, Armstrong Co. ; Walnut Bend, Venango Co.; Tionesta and Hickory, Forest
Co.; Warren, Warren Co.; Larabee, McKean Co. (Dennis Dally).

Buffalo Creek, Harbison, Butler Co.

Conemaugh River, New Florence, Westmoreland Co.

Loyalhanna River, Idlepark and Ligonier, Westmoreland Co.

Quemahoning Creek, Stanton’s Mill, Somerset Co.

Crooked Creek, Rosston and South Bend, Armstrong Co.

Little Mahoning Creek, Goodville, Indiana Co.

French Creek, Cochranton, Meadville, and Cambridge Springs, Crawford Co.

Leboeuf Creek, Waterford, Erie Co.

Connewango Creek, Russell, Warren Co.

M onongahcla-drainage :

Monongahela River, Elizabeth, Allegheny Co. (D. A. Atkinson).

Tenmile Creek, Clarksville, Greene Co.

South Fork of Tenmile Creek, Waynesburg, Greene Co.

Dunkard Creek, Wiley and IMount Morris, Greene Co.

Cheat River, Cheat Haven, Fayette Co.

Lake Erie-drainage:

Conneaut Creek, AVest Springfield, Erie Co.; Springboro, Crawford Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Genesee River, Rochester, Monroe Co., New York (R. H. Santens).

Chagrin River, Cuyahoga Co., Ohio (Hartman collection).

Sandusky River, Upper Sandusky, Wyandot Co., Ohio (C. Goodrich).

Maumee River, Roche de Boeuf Rapids, Lucas Co.; Defiance, Defiance Co., Ohio (C. Goodrich).
Beaver Creek, AVilliams Co., Ohio (C. Goodrich).

Raisin River, Adrian and Tecumseh, Lenawee Co., Michigan (C. Goodrich).

ORTMANN: monograph of the naiades of PENNSYLVANIA.

305

Ohio-drainage:

Lake Chautauqua, Chautauqua Co., New York (D. R. Sumstine; P. E. Nordgren; Miss B. Ortmann).
Tuscarawas River, Ohio (Holland collection).

Scioto River, Kenton, Hardin Co., Ohio (C. Goodrich).

West Fork White River, Riverside, Greene Co., Indiana (J. D. Haseman).

Wabash River, Bluffton, Wells Co., Indiana (C. Goodrich).

Ohio River, Toronto, Jefferson Co., Ohio; Portland, Meigs Co., Ohio.

Cheat River, Jaco and Mont Chateau, Monongalia Co., West Virginia.

West Fork River, Lynch Mines, Harrison Co.; West Milford, Harrison Co. (W. F. Graham); Lightburn
and Weston, Lewis Co., West Virginia.

Little Kanawha River, Grantsville, Calhoun Co. (W. F. Graham); Burnsville, Braxton Co., West Virginia.
North Fork Hughes River, Cornwallis, Ritchie Co.; Harrisville, Ritchie Co. (W. F. Graham), West
Virginia.

Pocatalico River, Raymond City, Putnam Co., West Virginia.

Elk River, Shelton and Clay, Clay Co.; Gassaway and Sutton, Braxton Co., West Virginia.

Little Coal River, Logan Co., West Virginia (Hartman collection).

Mud River, Milton, Cabell Co., West Virginia.

Levisa Fork Big Sandy River, Prestonsburg, Floyd Co., Kentucky.

Licking River, Farmer, Rowan Co., Kentucky.

Tennessee-drainage :

Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).

Shoals Creek, Lauderdale Co., Alabama (H. H. Smith).

Elk River, Fayetteville, Lincoln Co., and Estill Springs, Franklin Co., Tennessee (H. H. Smith).
Hurricane Creek, Gurley, Madison Co., Alabama (H. E. Wheeler).

Paint Rock River, Trenton, Jackson Co., Alabama (H. H. Smith).

Little River, Melrose, Blount Co., Tennessee.

Nolichucky River, Chunns Shoals, Hamblen Co., Tennessee.

Little Pigeon River, Sevierville, Sevier Co., Tennessee.

Holston River, McMillan and Mascot, Knox Co.; Hodges, Jefferson Co.; Turley Mill, Noeton, and
Holston Station, Grainger Co.; Austin Mill, Hawkins Co., Tennessee.

South Fork Holston River, Pactolus, Bluff City, and Emmett, Sullivan Co., Tennessee.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia; Mendota,
Washington Co., Virginia; Saltville, Smyth Co., Virginia.

Big Mocassin Creek, Mocassin Gap, Scott Co., Virginia.

Clinch River, Solway, Knox Co.; Edgemoor and Clinton, Anderson 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; Raven, Richland, and Cedar
Bluff, Tazewell Co., Virginia.

Emory River, Harriman, Roane Co., Tennessee.

Powell River, Combs, Claiborne Co., Tennessee; Dryden, Lee Co., Virginia.

Mississippi-drainage and westward:

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection); Moline, Rock Island Co .,
Illinois (P. E. Nordgren).

James River, Galena, Stone Co., Missouri (A. A. Hinkley).

White River, Cotter and Norfolk, Baxter Co., Arkansas (A. A. Hinkley).

306

MEMOIRS OF THE CARNEGIE MUSEUM.

Black River (H. E. Wheeler), and Spring River (A. A. Hinkley), Black Rock, Lawi’ence Co., Arkansas.
Illinois River, Tahlequah, Cherokee Co., Oklahoma (F. B. Isely).

Fourteen Mile Creek, Fort Gibson, Muskogee Co., Oklahoma (F. B. Isely).

Distribution and Ecology in Pennsylvania (See fig. 32) : This is a very common
form in western Pennsylvania, and is practically found everywhere, both in large
rivers and in small creeks. It is often associated with the typical L. ovata, in the
large rivers, but is distinctly less abundant in these, and, as has been said, inter-
grades with L. ovata. In the smaller streams where typical ovata is missing it is
extremely abundant, and becomes at favorable localities the prevailing species.
It goes far uj) into the headwaters, eastward to IMcKean, Indiana, Westmoreland,
and Somerset Counties. It should be noted, that it is also found in Gonneaut
Creek in the Lake Erie-drainage.

In its ecological preferences this form differs decidedly from the normal
L. ovata, for it does not favor rough bottoms and currents, but rather quiet pools
and eddies above and below riffles. Oiien ]iools in riffles, in Dianther a-psLiches,
with moderate current, and a liottom of fine gravel covered with a thin layer of
mud, seem to bo most favorable habitats, and occasionally this form is found
even in pure sand and rather deep and soft mud. It is the variety of smaller
streams, where it is found in rather protected locations, while L. ovata is the variety
of the rough waters of larger streams.

General distribution: Type locality, Wisconsin River (Barnes).

L. ovata vcntricosa is found all over the Ohio and upper Mississippi drainages
in western Pennsylvania, West Virginia, Ohio (Sterki, 1907a), Indiana (Call,
1896a & 1900), Illinois (Baker, 1906; Forbes & Richardson, 1913), Iowa (Pratt,
1876; Witter, 1878; Geiser, 1910), Wisconsin (Barnes, 1823; Cooper, 1855), and
Minnesota (Cooper, 1855; Grant, 1886; Holzinger, 1888). It also largely crosses
over into the lake-drainage in northern Illinois, Indiana, and Ohio, and occurs in
IMichigan (Walker, 1898). In New York, it is found in the upper Allegheny-
drainage (Chautauqua Lake), and at many places in the St. Lawrence-drainage
(Marshall, 1895). ‘ That the normal form is found here is shown by specimens
from the Genesee River in the Carnegie Museum. In addition it has been reported
from the lower St. Lawrence system and its lakes (Erie and Ontario) ; but in this
region it is generally supplanted by the var. canadensis (See below), and since
this has not been separated from ventricosa by most previous authors, the mutual
geograiihical relation of these two forms is obscure. No localities from the Atlantic
drainage in central New York are known, and it is missing on the Atlantic slope,
with the exception of the upper Potomac River, where it has been introduced

ORTMANN: monograph op the naiades of PENNSYLVANIA. 307

artificially and accidentally in recent times (see Ortmann, Nautilus 26, 1912, p. 51,
and 1913a, p. 318, and Marshall, Nautilus 31, 1917, p. 40). On the other hand,
L. ovata ventricosa is reported to cross over, in the northwest, into the Red River
and Nelson River systems in Canada, but little is known about the form of this
region (See below, under canadensis).

In a southerly direction, records from Kentucky are scarce, and Wilson &
Clark (1914) report it only from two tributaries of the Cumberland (Harpeth &
Stones R.) in Tennessee. But it certainly is found in the upper Tennessee-drainage,
where it has a similar relation to L. ovata as elsewhere, being preeminently found
in smaller streams and in the headwaters.

Westward it goes through Missouri (Utterback, 1916) to southeastern Nebraska
(Try on, 1868; Call, 1885) and eastern Kansas (Scammon, 1906), and southwest-
ward, it is found in Arkansas and Oklahoma. But in this region it passes into the
southwestern race called satura Lea (Scammon, 1906). In the Alabama-drainage
it is represented by allied forms, which require closer investigation.

Baker (1898) and Scammon (1906) say that it is generally found on muddy
bottoms, and with some qualification this agrees with my observations in Penn-
sylvania.

Lampsilis ovata canadensis (Lea) (1857).

Lampsilis ventricosa canadensis (Lea) Walker, 1913, p. 21; Lampsilis ventricosa

lurida Simpson, 1914, p. 41.

Plate XIX, figs. 4, 5. *

Records from Pennsylvania :

Ortmann, 19096, p. 202 (as the lake-form of ventricosa).

Characters of the variety: This form differs from L. ovata ventricosa, of which
it is a local race, by its smaller size, and very light-colored epidermis, which is
light-yellow, or light grayish or greenish yellow, with or without greenish rays.
In some specimens, the color of the epidermis inclines toward reddish brown or
chestnut, chiefly so near the beaks.

In other respects this form agrees with L. ovata ventricosa, but old specimens
of the male sex are often more elongated than the average ventricosa (slightly

over one-and-one-half times as long as high).

L. H. D.

Size: (Males) 1. Erie, Cat. No. 61.3989 101 mm. 66 mm. 46 mm.

2. do. “ “ 61.3982 93 “ 60 “ 42 “

3. do. “ “ 61.4883 68 “ 46 “ 33 “

(Females) 4. Erie, Cat. No. 61.3982 (gravid) 97 “ 68 “ 45 “

5. do. “ “ 61.4884 83 “ 63 “ 40 “

6. do. “ “ 61.4882 72 “ 54 “ 40 “

308

MEMOIRS OF THE CARNEGIE MUSEUM.

Soft parts identical with those of L. ovata and L. ovata ventricosa. Glochidia
not observed.

Breeding season: Gravid females have been seen only on May 21 and 22, 1909.
Most of my specimens were collected in the month of July, and no gravid females
were among them. Thus the interim seems to be in this month.

Remarks: Simpson (1914) calls this form var. lurida (of ventricosa)^ and
believes that U. canadensis Lea is a synonym of ventricosa. However, I follow
Walker (1913) in considering the lake-form to be L. canadensis.

This form has the same relation to L. ovata ventricosa, as has L. luteola rosacea
to L. luteola. It is a small, stunted, light-colored lake-form, which under certain
conditions shows the same tendency to become brownish near the beaks. The
shell is very variable in color. Specimens from the surf-beaten beaches are very
light in color, pale yellow; while those from protected locations in Presque Isle
Bay are darker, and tend to have more or less chestnut color toward the beaks.
The rays also vary greatly, and often are entirely absent.

Old shells frequently show a peculiar variation in shape, not found in L.
ovata ventricosa, having a tendency to become more elongated. In this form the
growth-rests are also often more regular and more distinct than in ventricosa, but
this is not always the case. This character is best exhibited in specimens from
deep water, and less so in specimens from the surf. The specimens from Ottawa
River do not show this. They are also darker grayish green than the average
specimens from Lake Erie.

Localities in Pennsylvania represented in the Carnegie Museum:

Lake Erie, Presque Isle Bay, and outer beach of Presque Isle, Erie; beach at Miles Grove, Erie Co.

Other localities represented in the Carnegie Museum:

Ottawa River, Ontario, Canada (B. Walker, donor).

Lake Erie, Vermilion, (C. Goodrich), and Cedar Point, Erie Co., Ohio (C. Brookover; O. E. Jennings);
La Plaisance Ba}", Monroe Co., Michigan (C. Goodrich); Port Rowan and Port Dover, Norfolk
Co., and Port Colborne, Welland Co., Ontario, Canada (C. Goodrich).

Lake Huron, Saginaw Bay, Charity Island, Michigan (C. Goodrich).

Distribution and Ecology (See fig. 32): Type locality, St. Lawrence River,
Montreal, Canada (Lea).

Besides the St. Lawrence, this form is found in Ottawa River, and very likely
at other localities in the lower St. Lawrence basin and Lake Ontario, generally
credited to L. ventricosa (Marshall, 1895; Whiteaves, 1863; Bell, 1859). It is

ORTMANN: monograph of the naiades of PENNSYLVANIA.

309

common in Lake Erie (Walker, 1913), and is also in Lake Huron. Whether it
extends farther west, remains to be seen.^^®

In the Lake Erie region L. ovata canadensis is restricted to the lake proper,
and does not go into the tributaries. In the lake this form is found in protected
bays as well as in the open lake, and it is one of the few species, which are found
in the surf, where I repeatedly picked up living specimens thrown out by the
waves. In Presque Isle Bay it is common everywhere, among rushes, and on
open shores, going down to a depth of about fifteen feet. It is here mostly on
sandy bottom. I never found it in the beach-pools.

Lampsilis fasciola Rafinesque (1820).

Lampsilis multiradiata (Lea) Simpson, 1914, p. 55.

Plate XX, figs. 1, 2.

Records from Pennsylvania :

Harn, 1891 (western Pennsylvania).

Marshall, 1895 (Allegheny River, Warren Co.)

Rhoads, 1899 (Beaver River, Wampum, Lawrence Co.)

Ortmann, 19096, p. 189.

Characters of the shell: The species is best described in terms of comparison
with L. ovata ventricosa. It is much smaller, and relatively a little thicker. It has
a more regularly ovate or elliptical outline. The posterior ridge is quite indistinct.
But the chief difference is in the color-pattern of the epidermis. This is yellowish
or greenish, or light brownish olive, with very numerous and crowded rays. The
latter may be rather broad, or they may consist of bundles of fine (capillary) rays,
and they are more or less wavy or interrupted. There are always many fine rays
between the wider ones, and often only fine rays are present. The interrupted and
wavy character is especially noticeable in the posterior part of the shell. Generally
the rays are distinct, but sometimes they are indistinct in the anterior part of the
shell. I have never seen a shell in which rays were entirely absent. According to
my experience, the nacre is always white.

I have two specimens (male and female) of a form of ventricosa from Sheyenne River, Argusville,
Cass Co., North Dakota (S. M. Edwards), which represent a depauperate form, but differ from L. cana-
densis by the more elongated shape and the more compressed valves. The material is too insufficient
to permit the expression of an opinion. L. ventricosa has been reported from the Red River of the North
and from Nelson River, but little is known about these forms.

According to Vanatta (1915, p. 552), Lampsilis fasciola of Rafinesque is L. luteola (Lamarck).
Nevertheless in this case I do not accept the determination of the so-called (but questionable) “ type,”
since Rafinesque’s description is unmistakable, and refers to an ovate (not elongate) shell, with wavy,
unequal rays. This fits only the present species.

310

MEMOIRS OF THE CARNEGIE MUSEUM.

The sexual differences agree with those of L. ovata ventricosa.

L. H. D.

Size: (Males) I. Shenango, Cat. No. 61.4099 88 mm. 58 mm. 39 mm.

2. do. “ “ do ■. ...62 “ 42 “ 29 “

3. do. “ “ do 44 “ 29 “ 18 “

(Females) 4. Russell, Cat. No. 61.3433 (gravid) 72 “ 53 “ 35 “

5. New Galilee, Cat. No. 61.3240 (gravid) 67 “ 51 “ 34 “

6. Wurtemberg, Cat. No. 61.4890 (gravid) 47 “ 36 “ 18 “

The largest male listed above is the largest individual I have ever seen.

Soft parts (See Ortmann, 1912, p. 352). Glochidia (See Lea, Obs. VI, 1858,
PI. 5, fig. 17; Ortmann, 1. c.; Surber, 1915, p. 1, PL 1, fig. 2). Surber gives the
following dimensions: 0.230 X 0.290 mm.; while Tgave: 0.25 X 0.29 mfn.

Breeding season: The dates for gravid females are rather complete, extending
from Aug. 4 to Oct. 23, and from IMay 14 to Aug. 9 (the last date furnished dis-
charging specimens) . Thus we would have the seasons overlapping early in August.
The species is clearly bradytictic. (Discharge has been repeatedly observed in
July.)

Remarks: This species is easily recognized and distinguished from L. ovata
ventricosa by its small size, regular outline, and the character of the rays. I never

• Lampsilis fasciola.

B Lampsilis orbiculata.

had any trouble in identifying it, and I have never seen any tendency to intergrade
with ventricosa, and consider it a well-marked species.

OETMANN: monograph of the naiades of PENNSYLVANIA.

311

Localities in Pennsylvania represented in the Carnegie Museum:

Oliio River, Industry, Beaver Co.

Little Beaver Creek, Cannelton (H. H. Smith), Darlington, and. New Galilee, Beaver Co.

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).

Connoquenessing Creek, Ellwood City, Lawrence Co. (G. L. Simpson, Jr.).

Slipperyrock Creek, Wurtemberg and Rose Point, Lawrence Co.

Mahoning River, Mahoningtown, Lawrence Co.

Neshannock Creek, Eastbrook, and Volant, Lawrence Co.; Leesburg, Mercer Co.

Shenango River, Pulaski, Lawrence Co.; Shenango, Mercer Co.

Pymatuning Creek, Pymatuning Township, Mercer Co.

Allegheny River, Aladdin, Godfrey, Johnetta, Kelly, and Templeton, Armstrong Co.; Walnut Bend,
Venango Co.; Tionesta and Hickory, Forest Co.; Warren, Warren Co.

Buffalo Creek, Harbison, Butler Co.

Conemaugh River, New Florence, Westmoreland Co.

Loyalhanna River, Idlepark, Westmoreland Co.

Quemahoning Creek, Stanton Mill, Somerset Co.

Crooked Creek, Rosston, Armstrong Co.

French Creek, Utica, Venango Co.; Cochranton, Crawford Co.

Connewango Creek, Russell, Warren Co.

Cheat River, Cheat Haven, Fayette Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Sandusky River, Upper Sandusky, Wyandot Co., Oliio (C. Goodrich).

Maumee River, Defiance, Defiance Co., Olno (C. Goodrich).

Raisin River, Tecumseh, Lenawee Co., and Grape P. 0., Monroe Co., Michigan (C. Goodrich).
Ohio-drainage :

White River, Rockford, Jackson Co., Indiana (G. H. Clapp, donor).

Cheat River, Jaco and Mont Chateau, Monongalia Co., West Virginia.

West Fork River, Lynch Mines, Harrison Co.; Lightburn and Weston, Lewis Co., West Virginia.

Little Kanawha River, Burnsvill, Braxton Co., West Virginia.

Elk River, Shelton, Clay Co.; Gassaway and Sutton, Braxton Co., West Virginia.

Coal River, Sproul, Kanawha Co., West Virginia.

Tennessee-drainage :

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).

Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).

Shoals Creek, Lauderdale Co., Alabama (H. H. Smith).

Blue Water Creek, Lauderdale Co., Alabama (H. H. Smith).

Elk River, Estill Springs, Franklin Co., Tennessee (H. H. Smith).

Flint River, Maysville and Gurley, Madison Co., Alabama (H. H. Smith).

Hurricane Creek, Gurley, Madison Co., Alabama (H. E. Wheeler).

Paint Rock River, Paint Rock, Trenton, and Princeton, Jackson Co., Alabama (H. H. Smith).

South Chickamauga Creek, Ringgold, Catoosa Co., Georgia.

Little River, Melrose, Blount Co., Tennessee.

312

MEMOIRS OF THE CARNEGIE MUSEUM,

Little Pigeon River, Sevierville, Sevier Co., Tennessee.

Pigeon River, Canton, Haywood Co., North Carolina.

liolston River, McMillan and Mascot, Knox Co.; Turley Mill and Holston Station, Grainger Co.,
Tennessee.

South Fork Holston River, Pactolus, Bluff City, and Emmett, Sullivan Co., Tennessee; Barron, Wash-
ington Co., Virginia.

Watauga River, Watauga, Carter Co., Tennessee.

Middle Fork Holston River, Chilhowie, Smyth Co., Virginia.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia; Mendota,
Washington Co., Virginia; Saltville, Smyth Co., Virginia.

Big Mocassin Creek, Mocassin Gap, Scott Co., Virginia.

Clinch River, Edgemoor and Clinton, Anderson Co.; Black Fox Ford, Union Co.; Clinch River Station,
Claiborne Co.; Oakman, Grainger Co., Tennessee; Clinchport, Scott Co., Virginia; St. Paul,
Wise Co., Virginia; Fink and Cleveland, Russell Co., Virginia; Raven, Richland, and Cedar Bluff,
Tazewell Co., Virginia.

Powell River, Combs, Claiborne Co., Tennessee; Dryden, Lee Co., Virginia.

Distribution and Ecology in Pennsylvania (See fig. 33) : This species has about
the same distribution in western Pennsylvania as L. ovata ventricosa, but it is missing
in the Lake Erie-drainage. In the Ohio-drainage it is practically generally dis-
tributed, but is cpiite scarce, and never found in great numbers. In the Ohio
below Pittsburgh I found it only once, and also in the Allegheny River in Arm-
strong County. Although found at a number of localities, it is rare. Farther
up in the Allegheny and its tributaries it is met with regularly. It is possibly
most abundant in Little Beaver Creek, and in the drainage of Beaver River. In
the whole Monongahela-drainage it has been found only in the Cheat River, and
more abundantly in West Virginia. It is to be noted that it goes up in the Cone-
maugh-drainage into the mountain-streams in Westmoreland and Somerset Coun-
ties.

In Pennsylvania it distinctly prefers riffles with lively currents and gravelly
bottoms, but it is not found in very strong currents and among large rocks. The
most favorable localities seem to be riffles with an abundant growth of Dianthera
amei'icana and other water weeds. It distinctly differs in its ecology from L.
ovata ventricosa, although often found in close proximity to it. But while the
latter prefers more quiet water above and below the riffles, L. fasciola is found in
the riffles.

General distribution: Type locality, Kentucky River (Rafinesque).

According to Simpson (1900) it is found in the entire Ohio-drainage; in
southern Michigan and in New York. This is essentially correct. In the Ohio-
drainage it is jiractically everywhere, in western Pennsylvania, West Virginia, in
Ohio (Sterki, 1907a), and Indiana (Call, 1896a, 1900). In Illinois it seems to be

ORTMANN: monograph of the naiades of PENNSYLVANIA.

313

less widely distributed, but has been reported from the southern as well as the
northern parts of the state (Baker, 1906). Records from Kentucky are scarce
(type locality and Green River, according to Call, 1885). In the Cumberland it
is rare in the main river, but common in the tributaries (Wilson & Clark, 1914).
In the Tennessee-drainage it seems to be abundant, from Tennessee and Alabama
up to the affluents in Virginia, being found in this region chiefly in the smaller
streams. In this region it has also been reported (by Call, 1885) from the French
Broad River in North Carolina, and I found it in the Pigeon River in North Caro-
lina.

South of the Tennessee and West of the Mississippi it is absent.

In addition it has crossed over into the lake-drainage in Indiana, St. Joseph
River, Michigan basin, and Maumee River (Call, 1896a) and Ohio (Sterki, 1907a),
and is found in the latter state and in southeastern Michigan in the drainage of
Lake Erie (Walker, 1898).^°^ It is not known to go up in the Allegheny-drainage
into New York state, but Marshall (1895) gives it from Medina, Orleans Co.,
and the Genesee River, Monroe County, in the Lake Ontario-drainage, and from
Butternut Creek in Otsego County (tributary to the Susquehanna) . These locali-
ties should be confirmed, and Simpson treats them as doubtful.

Baker (1898a) says that, like Eurynia iris, this species is found in lakes and
rivers on a sandy or muddy bottom. This is not at all confirmed by my observa-
tions in Pennsylvania. In the upper Tennessee-drainage, where the species is
abundant, it prefers rather rough parts of the streams, and ascends (in French
Broad and Pigeon Rivers) into the high Appalachians in North Carolina.

Lampsilis cariosa (Say) (1817).

Lampsilis cariosa (Say) Simpson), 1914, p. 43.^“®

Plate XX, figs. 3, 4, 5.

Records from Pennsylvania:

Say, 1817 (Delaware and Schuylkill Rivers, Philadelphia).

Haldeman, 1844 (Lancaster Co.).

201 Maumee River (Dali & Simpson, 1895) and Cuyahoga River (Dean, 1890) in Ohio, and Huron
and Detroit Rivers, in Michigan (Walker, 1892, see also material in Carnegie Museum); also reported
from Lake Erie proper by Walker, 1913, but never found on the Pennsylvanian shores.

202 Not 1816.

203 The figures of Conrad (Mon. 4, 1836, PI. 19) are fine representations of this species, and the
same is true of his figures of U. ochraceus (PI. 17, fig. 2). If subsequent authors had paid due attention
to these figures, the prevailing confusion of these two species (and of the related western forms) would
have been impossible. DeKay s figures (1843, PI. 21, figs. 243, 244) surely do not represent cariosus,
but are L. ovata ventricosa. Simpson (1895) on pp. 121 and 122 also figures cariosus and ochraceus, and

I

314 MEMOIRS OF THE CARNEGIE MUSEUM.

Gabb, 1861 (Delaware River, Tacony, Philadelphia Co., and Bristol, Bucks Co.)

Bruckhart, 1869 (Lancaster Co.)

Hartman & Michener, 1874 (Schuylkill, Delaware, and Susquehanna Rivers).

Pilsbry, 1894 (York Furnace, York Co.)

Schick, 1895 (Delaware River, Philadelpliia).

Ortmann, 19096, p. 204.

Caffrey, 1911, reports U. ochraceus from the Delaware in Northampton County, but there is no question
that under this name he designates U. cariosus.

Characters of the shell: Of medium size, rather thin when young, but becoming
thicker when old. Outline ovate or subelliptical, rather short and high, but old
males are often somewhat elongated, and may be slightly over one-and-a-half
times as long as high; but generally the length is less than this. Anterior margin
rounded. Lower margin more or less curved. Upper margin short, straight or a
little convex, passing into a blunt angle or gradually into the obliquely descending-
posterior margin. Posterior end a rounded, blunt angle. Beaks moderately
swollen, not much elevated, anterior to the middle of the shell. Beak-sculpture
(Marshall, 1890, fig. 8) consisting of about five not very distinct bars, the first
subconcentric, the following ones slightly double-looped, with a light sinus in the
middle, obsolete in the last bars. Upon the posterior slope, the bars are effaced.
Valves regularly convex, somewhat flattened upon the sides. Posterior ridge very
indistinct, practically absent. Posterior slope gently convex, or almost flat.

Epidermis yellow, very smooth and shining. The yellow is wax- or straw-
yellow, with hardly a trace of green in it, but sometimes inclining to brown or
reddish brown. Rays either entirely absent, or present upon the posterior slope
and a little in front of it. They are variable in width, but are generally fine,
straight, and sharply defined, dark green or blackish in color, contrasting strongly
with the light epidermis. Concentric bands of light or dark color absent or in-
distinct. In old specimens the color of the epidermis is less bright, and becomes
dirty grayish or brownish yellow.

Hinge well-developed. Pseudocardinals generally two in each valve, quite
variable in shape. In young specimens they are rather compressed, in older ones
more stumpy and ragged. Interdentum practically absent. Laterals lamellar,
generally somewhat elevated and truncate at their posterior ends. Beak-cavity
moderate. Dorsal muscle-scars in the beak-cavity. Anterior adductor-scars
distinct and impressed, posterior ones less distinct. Nacre silvery white, often
suffused with cream- or salmon-color, but not reddish.

later (1900, p. 529, footnote 1, and p. 530, footnote 2) says, that the two figures had been accidentally
transposed. But these figures are copies from Hartman & Michener (1874, figs. 183, 184) where the
same mistake is made. I think, both of these figures represent cariosus, and the one named ochraceus
is certainly the female of cariosus.

ORTMANN: monograph of the naiades op PENNSYLVANIA.

315

Sexual differences of shell strongly marked. In the male, the lower margin
is more regularly convex, joining the posterior margin in a blunt posterior point,
and the outline of the shell is more regularly elliptical. In the female, the lower
margin is considerably expanded in the posterior region, sloping up very strongly
to the blunt posterior end, so that the posterior part of the shell is higher and more

broadly rounded.

L. H. D.

Size: (Males) 1. Selinsgrove, Cat. No. 61.4894 122 mm. 80 mm. 51 mm.

2. Duncannon, Cat. No. 61.3416 113 “ 74 “ 41 “

3. Yardley, Cat. No. 61.3417 80 “ 50 “ 32 “

(Females) 4. York Haven, Cat. No. 61.4893 96 “ 71 “ 44 “

5. South Waverly, Cat. No. 61.4003 (gravid) 91 “ 67 “ 46 “

6. Yardley, Cat. No. 61.3417 (gravid) 70 “ 52 “ 36 “

Soft parts (See Ortmann, 1912, p. 352).' Glochidia, ibid, (the latter not fully
known, only immature individuals having been observed) .

Breeding season: Gravid females were observed on the following dates; Aug.
13, 1908; Aug. 14, 1908; Aug. 14, 1910; Aug. 20, 1909; Aug. 22, 1909; Aug. 24,
1908. Only on the last date were glochidia (immature) found; all other specimens
had only eggs. But just these meagre facts establish the beginning of the breeding
season in August, and very probably this species agrees with the allied forms, and
is hradytictic, discharging the glochidia in the subsequent spring or early summer.

Remarks: An exceedingly weU defined species, easily recognized. Neverthe-
less great confusion prevails with regard to it. This confusion is principally due
to the fact that De Kay (1843) misunderstood this and some allied species, and
that the two sexes of cariosus were regarded as different species by other authors
(Gould-Binney, and Hartman & Michener). Simpson (1895) tried to straighten
out the confusion, but unfortunately copied two old erroneous figures, and com-
mitted an additional error, and thus it has happened that even some recent authors
(Conner, 1909, and Caffrey, 1911) confused this species with L. ochracea.

L. cariosa is allied to the western L. ovata ventricosa, but is smaller than the
latter, in the male sex is more regularly elliptical, has hardly a trace of a posterior
ridge, and is entirely and characteristically different in color and color-markings.
The bright and light yellow of the epidermis and its high gloss are entirely different
from the greenish-olive tints of L. ovata ventricosa, and the restriction of the rays
(if such are present) to the posterior section of the shell is also characteristic. Of
course, these characters are best seen in young or medium-sized individuals, while
in older specimens the smoothness disappears, and the color becomes more dirty:
n evertheless even in very old shells the light yellow is generally present at least on

316

MEMOIRS OF THE CARNEGIE MUSEUM.

certain parts of the shell. This color is so eminently characteristic, that even
laymen have noticed it, and people living on the banks of the Susquehanna and
Delaware properly distinguish this species as the “yellow clam.”

The variation of this species occurs chiefly in the external shape (more or less
elongate) and the development of the rays. Moreover the yellow of the ground-
color varies slightly, being lighter or more intense, often inclining toward reddish
brown in parts of the shell.

Localities in Pennsylvania represented in the Carnegie Museum:

Delaware River, Taylorsville (C. H. Conner) and Yardley, Bucks Co.; Shawnee, Monroe Co.^°^
Susquehanna River, York Furnace and York Plaven, York Co.; Duncannon, Perry Co.; Selinsgrove,
Snyder Co.

Conewago Creek, York Haven, York Co.

Juniata River, Juniata Bridge, Perry Co.; Lewistown, Mifflin Co.

Frankstown Branch, Juniata River, Huntingdon, Huntingdon Co. (D. A. Atkinson).

Raystown Branch, Juniata River, Ardenheiin, Huntingdon Co.

West Branch Susquehanna River, Williamsport, Lycoming Co. (D. A. Atkinson).

North Branch Susquehanna River, Tunkhannock, Wyoming Co.

Chemung River, South Waverly, Bradford Co.

Other localities represented in the Carnegie Museum:

Connecticut River (Hartman collection).

Delaware River, Newbold, Gloucester Co., New Jersey (C. H. Conner).

Potomac River, Cabin John, Montgomery Co., Maryland (J. D. Hasemqn); Mount Vernon, Fairfax
Co., Virginia (Juny collection).

Savannah River (Hartman collection).^®®

Distribution and Ecology in Pennsylvania (See fig. 31): This species belongs
to the Delaware and Susquehanna drainages in Pennsylvania, but has not been
found in the small tributaries of the Potomac in our state. In the Delaware River
it goes up to above the Delaware Water Gap, but its scarcity just above the gap
indicates, that at this point it approaches the upper limit of its range. In the
Susquehanna it is practically everywhere, ascending at least to Williamsport,
Lycoming Co., in the West branch, and to the New York state line in the North

Shawnee is immediately above the Delaware Water Gap: only one specimen was found here and
no dead shells were lying around.

In the lowermost part only of this creek, where there is at times water from the Susquehanna.
A large, thick-shelled specimen, labeled rosaceus Conrad, agrees well with Conrad’s description,
but is larger, and has only a faint blush of pink on the nacre. In other respects it agrees likewise with
cariosus, and, although discolored, the epidermis shows remnants of the gloss. If locality and identifica-
tion are correct, rosaceus is a form of cariosus, not of ochraceus (of which Simpson makes it a synonym).
Whether this form is different from cariosus (as variety or species) cannot be decided from a single indi-
vidual.

OETMANN: MONOGEAPH of the naiades of PENNSYLVANIA.

317

branch. “It is generally restricted to the main river, and only enters some of the
larger tributaries, for instance the Schuylkill, Juniata, and Chemung Rivers.
In the smaller tributaries it is absent, and the only specimen found by myself in
Conewago Creek, was in its lowermost part, which, during high water forms a
branch of the Susquehanna.

Where found this species is generally abundant, even in smaller streams like
the Juniata in Huntingdon County, where it reaches its farthest advance in the
mountains. It is always found in lively currents, on shoals and riffles, in finer or
coarser gravel, and very often in bars of pure sand. Although it has been reported
from the tidewater region of the lower Delaware, it does not seem to be abundant
there (I did not find it near Fish House, Camden Co., New Jersey).

In Pennsylvania its metropolis is apparently the Delaware from Trenton
upward, and the Susquehanna from the region of York Haven to the point where
the West and North branches unite.

General distribution: Type locality, Delaware River, Philadelphia (Say).

According to Simpson (1900) this species is found in the “ Atlantic-drainage
from Georgia to the lower St. Lawrence,” but this possibly requires some restric-
tion. Its occurrence in the lower St. Lawrence is extremely questionable, and
rests chiefly upon the statement of Marshall (1895), who also gives it from Maine
(confirmed by Lermond, 1909, and Johnson, 1915). It is undoubtedly found in
Massachusetts and Connecticut (Gould-Binney, 1870, Marshall, 1895, Linsley,
1848, Johnson, 1915). From New York it has been reported (Marshall, 1895)
from the Hudson-drainage up to, and even above, Albany, and there seems to be
even a western extension of the range through the Erie canal to Onondago and
Ontario Counties, but this should be investigated again, since it is not quite sure
that Marshall understood this species correctly. De Kay (1843) reports it from
the Passaic River, Belleville, Essex Co., New Jersey, and the species should exist
there: however, De Kay’s figures represent, as has been said, L. ovata ventricosa.
Marshall (1895) gives it from the Raritan River, Somerville, Somerset Co., New
Jersey. Its distribution in Pennsylvania has been discussed above. From Dela-
ware it is known from Seaford, Sussex Co. (Rhoads, 1904). From the Potomac
River it has been reported from the District of Columbia and from the canal at
Alexandria, Fairfax Co., Virginia (Marshall, 1895), and it is found in Maryland
above Washington. However, it does not ascend far up in this river, and is surely
absent in the whole Potomac-drainage West of the Blue Ridge.

To the South of the Potomac, records become very scarce. Conrad (1836)
says that it is rare in the Potomac as well as in the James River, and since Conrad

318

MEMOIRS OP THE CARNEGIE MUSEUM.

correctly understood the species, we must accept this latter locality, “Georgia,”
given by Simpson, probably rests upon Conrad’s record of U. oratns from Flint
River, and if U. rosaceus Conrad should actually belong here, we would also have
to include the Savannah River, However these southern localities emphatically
need confirmation in view of the fact that from the James River southward, in
Virginia and the Carolinas, not a single reliable record is at hand.

As to the standing of this species in the Atlantic fauna, see Ortmann, 1913a,
pp. 325 & 363.

Lampsilis ochracea (Say) (1817).^'^^

L. ochracea Simpson, 1914, p. 49.^°*

Plate XX, figs. 6, 7.

R ecords f rom Pennsylvania :

Say, 1817 (Delaware and Scliuylkill Rivers, “ with the preceding,” namely TJ. cariosus).

Conrad, 1836 (Dela.ware and Schujdkill Rivers).

Gabb, 1861 (Schuylkill and Wissahickon, Philadelphia; League Island, Philadelphia).

Schick, 1895 (Dejaware River, Philadelphia).

IVIarshall, 1895 (Philadelphia).

Hartman & Michener have confounded this species with L. cariosa, and their records: Schuylkill,
Delaware, and Susquehanna are unreliable. This species has never been found in the Susquehanna in
Pennsylvania. Caffrey (1911) records it from the Delaware, Northampton Co., but he certainly refers
to L. cariosa. See also Ortmann (19096, pp. 204 & 209).

Characters of the shell: Having only scanty material at hand, I confine myself
to giving only the characters of this species differentiating it from its allies, chiefly
L. cariosa.

L. ochracea is a rather small, thin shell, much thinner than L. cariosa, and
much thinner and smaller than any of its M^estern relations {ventricosa, fasciola).
It resembles these in outline, but is mostly somewhat shorter than L. cariosa,
and has a slightly more distinct posterior ridge. The chief difference is in the
epidermis, which does not have the gloss of L. cariosa, and becomes quite rough
toward and on the posterior slope. The color is dull, not bright yellow, but gray-
ish, greenish, yellowish, or brownish olive, a grayish green being the prevailing
hue; and the rays have a different character. Sometimes the latter are entirely
absent, but (when present) they are not sharp and blackish, but indistinct, and
grayish or grayish green, rather fine, and cover all or a large portion of the surface.
Upon the posterior slope, the rays are obscure, and sometimes somewhat wider.

2°^ Not 1816.

The figures of DeKay (1843, PI. 19) do not represent this species, but L. radiata (See above, p.
292, footnote 191). Very probably the figure of Goidd-Binney (1870, p. 174, fig. 476), and certainly that
of Hartman & Michener (1874, p. 89, fig. 184) represent females of L. cariosa. There is only one good
figure of this species, that of Conrad (Mon. 4, 1836, PI. 17, fig. 2) which represents the male.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

319

The nacre in all my specimens is silvery white, but is said to be sometimes more or
less reddish. Corresponding to the thinness of the shell, the hinge-teeth are deli-
cate, thin, and compressed. The sexual differences of the shell are similar to those
of L. cariosa. Beak-sculpture as described by Marshall (1890, fig. 7), and re-
sembling that of L. cariosa^ but the median sinus of the bars is very indistinct or
even missing. I have only a single individual which shows the beak-sculpture.

L. H. D.

Size: I. Newbold, Cat. No. 61.4012 (cT) 62 mm. 42 mm. 26 mm.

2. do. “ “ do. (9) 46 “ 30 “ 20 “

3. Plymouth, Cat. No. 61.667 (9) 38 “ 29 “ 18 “

4. Princeton, Cat. No. 61.1571 (sex ?) 32 “ 22 “ 13 “

This species grows somewhat larger, but the largest male given above is about
the average size.

Soft parts and glochidia practically unknown. Lea (Obs. II, 1838, PL 15,
fig. 44) has given a poor figure of the soft parts of a gravid female, and an incom-
plete description (Obs. X, 1863, p. 455).

Breeding season: Through Lea (Obs. II, 1838, p. 54) we know that this species
is gravid in autumn (October, November). Lea also reports {1. c., p. 57) a female
charged with eggs for June 5. Conner (1909) has confounded this species with L.
cariosa, and thus we cannot depend on his dates.

Re7narks: This species has been often misunderstood and confounded with L.
cariosa. In general terms it. may be defined as a rather small and thin shell, with
dull-colored epidermis, and fine, indistinct, or missing rays, which, when present,
cover a larger part of the shell than is the case in L. cariosa.

Localities represented in the Carnegie Museum:

Plymouth, Plymouth Co., Massachusetts (Hartman collection).

Delaware-Raritan Canai, Princeton, Mercer Co., New Jersey.

Delaware River, Newbold, Gloucester Co., New Jersey (C. H. Conner) (received as L. cariosa).

Lake Wacamaw, Columbus Co., North Carolina (G. H. Clapp, donor).

Localities represented in the Philadelphia 'Academy.

Ditches of Delaware Meadows, League Island, Philadelphia, Pa.

Delaware River, Westville, Gloucester Co., New Jersey (Morris Schick).

Delaware River, Kaighns Point, Camden, Camden Co., New Jersey (John Ford).

Distribution and Ecology (See fig. 31) : Type locality, Delaware River, Phila-
delphia (Say).

According to Simpson (1900) this species is in the “ Atlantic-drainage, from
New England to the Ogeechee River, Georgia,’’ that is to say, it occupies about

320

MEMOIRS OF THE CARNEGIE MUSEUM.

the same range, as that given by him for L. cariosa. But, as we have seen, in
the south L. cariosa is doubtful.

The species has been reported from Maine (Marshall, 1895; Lermond, 1909),
from Massachusetts (Earle, 1835; Gould-Binney, 1870; Marshall, 1895), from
Connecticut (Linsley, 1845) (See also New England localities given by Johnson,
1915). It is known from the Hudson in New York, and is said to go up here to
Albany and the Mohawk River, Herkimer Co. (Marshall). In Pennsylvania, it
is restricted to the lower Delaware, and the lowlands near to it. It has been found
in the lower Schuylkill and also Wissahickon Creek. According to specimens I
have seen in the Philadelphia Academy, it is abundant in the ditches of the meadows
near League Island. It is unknown from the Susquehanna and Potomac drainages
in Pennsylvania, although known from the lower Potomac in the District of Colum-
bia (Marshall), Prince Ceorge Co., Maryland (Lea, Obs. X, 1863, p. 456), and
Fairfax Co., Virginia (Marshall). Rhoads (1904) reports it from Seaford, Sussex
Co., Delaware, and farther south it is abundant in the lower James River, Virginia
(Conrad, 1836) and according to the same author in “most tide-waters north of
the Savannah River.” The most southern record is that of Simpson from the
Ogeechee River in Georgia.

In Pennsylvania, L. ochracea is decidedly rare, and restricted to the tide-
waters of the Delaware. It is not at all coextensive with L. cariosa, since the
latter ascends the rivers for a considerable distance. The same seems to be true
in other parts. Most of the exact localities known are in the tidewater-regions
of the Atlantic streams, and Conrad calls especial attention to this. Thus the
ecological preferences of this species seem to be for the estuaries of our large rivers.
In the Hudson it goes up quite far, but in the vicinity of Albany it has been reported
chiefly from canals, and it seems that it also frequently inhabits canals and ponds.
I found it myself in the Delaware-Raritan Canal in New Jersey. Thus the species
differs ecologically from L. cariosa. It is a form of estuaries, ponds, canals, and
ditches, probably with more or less muddy bottoms, while L. cariosa favors large
rivers, with sandy and gravelly bottoms and strong currents. A closer study of
these conditions is very desirable (See Ortmann, 1913a, pp. 325, 368).

Lampsilis orbiculata (Hildreth) (1828).

Lampsilis orbiculata (Hildreth) Simpson, 1914, p. 76.

Plate XX, fig. 8; Plate XXI, figs. 1, 2.

Records from Pennsylvania:

Clapp, 1895 (Allegheny Co.)

Ortmann, 19096, p. 190.

ORTMANN; monograph of the naiades of PENNSYLVANIA.

321

Characters of the shell: Of medium size to rather large (but not as large as
L. ovata), extremely thick and heavy (one of the heaviest shells for its size). Out-
line subelliptical or subovate, very variable and often irregular, chiefly so in the
female, always less than one-and-a-half times as long as high. Anterior margin
rounded. Lower margin convex, sometimes rather uniformly so, but in other
cases the middle part of the lower margin is less convex, and the posterior part
slopes up rather steeply. Upper margin short, gently convex, passing gradually
into the obliquely descending posterior margin, without forming a distinct angle.
Posterior and lower margins uniting posteriorly in a blunt angle, which may be
obliterated. Beaks moderately swollen, and little projecting above the hinge-
line, more or less inclined forwards, and located in front of the middle of the shell.
Beak-sculpture obsolete. Although I possess specimens with fairly well preserved
beaks, I cannot see any distinct beak-sculpture. Valves more or less convex,
generally moderately so, but old specimens, chiefly females, are much more convex
(almost globular). The convexity is least upon the sides, greatest in the region
of the posterior ridge. The latter, however, is quite indistinct. Posterior slope
slightly convex or flattened, narrow.

Epidermis brown, light or dark yellowish brown or reddish brown, generally
without any trace of green shades. This color is rather uniform, and no concentric
bands of light and dark are present. Rays either absent (chiefly in older shells'),
or, when present, faint and indistinct. In young, well-preserved specimens, the
rays are fine or moderately wide, but not dark (black or dark green), but gray or
grayish green, or brownish. They are not sharply contrasted with the ground-
color, and disappear toward the lower margin, so that older shells have rays only
near the beaks. In very old specimens the general color becomes uniformly dirty
brown, but not very dark brown.

Hinge well developed. Pseudocardinals two in left, one or two in right valve
(the anterior one smaller), variable in shape, strong, stumpy and ragged. Inter-
dentum narrow. Laterals moderately long, slightly curved, thick and heavy.
Beak-cavity moderate. Dorsal muscle-scars in the beak-cavity. Adductor-scars
distinct, aU deeply impressed, the posterior scars less so. Nacre silvery white,
very often with more or less pink, chiefly in the beak-cavity and near the hinge,
and sometimes the whole interior of the shell is suffused with beautiful salmon or
pink. But in other cases the pink shades may be entirely absent.

Sexual differences of the shell very strongly marked. In the male the out-
line is rather regularly subovate or subelliptical, with the lower margin rather
regularly curved, meeting the posterior margin in a blunt point. In older males

322

MEMOIRS OF THE CARNEGIE MUSEUM.

the lower margin may be almost straight in the middle, and the posterior end may
be rounded off. In the female the lower margin projects in the postbasal region,
so that the anterior and middle part is almost straight, and the posterior part
ascends very suddenly, forming with the part in front a more or less distinct rounded
angle, and joining the posterior margin in a very blunt angle. Thus the posterior
part of the female shell becomes very high, and in old females this may go to such
an extreme that the height of the shell nearly equals the length, thus rendering the
outline almost orbicular. In some cases such old females are also much more
convex, so that the whole shell approaches the globular shape.

L. H. D.

Size: (Males) I. Portsmouth, Cat. No. 61.4898 103 mm. 75 mm. 61 mm.

2. Neville Island, Cat. No. 61.2046 100 “ 76 “ 57 “

3. Kelly, Cat. No. 61.3017 80 “ 59 “ 36 “

(Females) 4. Industry, Cat. No. 61.3500 (gravid) 92 “ 83 “ 57 "

5. do. “ “ do 89 “ 70 “ 49 “

6. Godfrey, Cat. No. 61.3504 74 “ 57 “ 34 “

The largest specimen I have seen is from the Clinch River, at Offut, Anderson
Co., Tennessee, and measures: L. 104, H. 82, D. 53 mm. It has the shape of the
female.

Soft 'parts (See Ortmann, 1912, p. 353). Glochidia (See Ortmann, 19116,
PI. 89, fig. 22).

Breeding season: The following records for gi’avid females are at hand. Aug.
10, 1909; Aug. 24, 1910; Aug. 29, 1908; Sept. 8, 1908; Sept. 10, 1908; Sept. 12,
1914; Sept. 17, 1908; Sept. 23, 1908. Sept. 8 is the earliest date for glochidia.
A female just discharged was collected in Ouachita River by Mr. Wheeler on June
26, 1911.

This is apparently a bradytictic form, beginning to breed in August, having
glochidia in September, which are discharged the next spring (June).

Remarks: This species, when well-developed, is easily recognized by the
extremely heavy shell, by the brown color with indistinct rays, and the general
presence of a delicate pink color in the nacre. However, it is very variable with
regard to shape. The males, and chiefly the young males, very much resemble
Actinonaias ligamentina, so much so in fact, that this species has been placed by
the side of L. ligamentina in Simpson’s system. Yet there is no close relationship
between the two species. Young males of L. orbiculata differ from A. ligamentina
by the brown color of the epidermis, and the indistinct and finer rays; however,
sometimes the latter species also has a brownish epidermis, and in such cases it is
hard to separate them.

ORTMANN: monograph of the naiades op PENNSYLVANIA.

323

The female of L. orhiculata is entirely different from that of A. ligamentina
in shape. The posterior dilatation is not found in A. ligamentina, and the latter
never approaches the globular shape.

Both male and female of L. orhiculata often present freakish shapes, and the
proportion of length and height, convexity of the valves, and curvature of the
lower margin is hardly alike in any two specimens.

A very good character, excepting in the case of very young shells, is always the
thickness of the shell. In fact for its size this species is possibly the heaviest shell
of the Ohio-drainage.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Shippingport, Cooks Ferry, and Industry, Beaver Co.; Neville Island, Allegheny Co.
Allegheny River, Godfrey and Kelly, Armstrong Co.

Monongahela River, Charleroi, Washington Co. (G. A. Ehrmann).

Other localities represented in the Carnegie Museum:

Ohio River, Toronto, Jefferson Co., Ohio; St. Marys, Pleasants Co., West Virginia; Portland, Meigs
Co., Ohio; Portsmouth, Scioto Co., Ohio.

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).

Clinch River, Solway, Knox Co., and Offutt, Anderson Co., Tennessee.

Mississippi River, Muscatine, Muscatine Co., Iowa (Hartman collection); Andalusia, Rock Island Co.,
Illinois (A. D. Howard).

Black River, Black Rock, Lawrence Co., Arkansas (H. E. Wheeler); Pocahontas, Randolph Co., Arkansas
(H. E. Wheeler).

Ouachita River, Arkadelphia, Clark Co., Arkansas (H. E. Wheeler)

Distribution and Ecology in Pennsylvania (See fig. 33) : This is a rare shell in
Pennsylvania, and is only found in the larger rivers. The largest number of speci-
mens has been secured in the Ohio in Beaver County, but here also it is by no
means abundant. It used to be in the Ohio in Allegheny County, and in the
Monongahela as far up as Charleroi, Washington Co. (only one individual is at
hand from this locality). It also ascends the Allegheny to Armstrong County,
but here it is extremely rare, and only a few specimens were taken.

So far as I am able to judge, it is a shell of strong currents in large rivers. I
found all my specimens on riffles; but farther down in the Ohio it is present upon

See Wheeler (1918, p. 117). Specimens from the Ouachita River agree very well in shape with
specimens from the Ohio River and the color of the epidermis is of the same brown hue in some of them,
but in others it is more brownish-olive (with a suggestion of green). They are surely not L. higginsi,
because the beaks are not at all inflated. The form from Black River is remarkable on account of its
rays, which are broad and distinct; and also by the brownish-olive epidermis. It belongs to the forms
intergrading with L. higginsi (Lea).

324

MEMOIRS OF THE CARNEGIE MUSEUM.

the shell-banks, among other bed-forming species. But here also it is not very
frequent, and the shell-heaps of the clam-diggers contain only small numbers.
These conditions remain the same all the way down to Portsmouth. Every
specimen found by myself has been taken, but the total sum collected from the
Ohio below Pennsylvania hardly amounts to two dozen.

General distribution: Type locality, Muskingum River, Marietta, Washington
Co., Ohio (Hildreth).

This species is found extending from western Pennsylvania down the Ohio
in the states of West Virginia, Ohio (Sterki, 1907a), Indiana (Call, 1896a), and
probably also Illinois and Kentucky. In the upper parts of the Ohio-drainage it
hardly goes into the tributaries. The type locality, Muskingum River, has re-
mained so far the only tributary, which contains it in Ohio. In Indiana it is also
found in the Wabash and White Rivers (Say, 1817, Call, 1896a and 1900), and in
Illinois (Baker, 1906) it is in the Wabash and in the Illinois River (as far up as
La SaUe Co.). It is in the Mississippi River in Illinois as well as in Iowa (Daven-
port, Scott Co.) according to Pratt (1876). Simpson (1900) says that it ascends
the Mississippi to Minnesota, but it has not been reported by Grant (1886) and
Holzinger (1888), and is not reported from Wisconsin by Lapham (1860).

From the southern tributaries of the Ohio it is known from the Cumberland
River (Call, 1885; Simpson, 1900; Wilson & Clark, 1914) and in the Tennessee, it
goes up to the Clinch River in Anderson Co., Tennessee.

West of the Mississippi previous records are scarce or doubtful. While Call
(1885) gives this species from Blue River in Kansas, Scammon (1906) records
L. higginsi for this river. The latter species, according to Simpson, is found
from the (probably lower) Ohio west to Iowa and southwest to Kansas.^”® How-
ever, specimens sent to me by H. E. Wheeler from Ouachita River in Arkansas are
undoubtedly L. orhiculata, and thus the latter certainly goes southward to the
Ouachita River. Specimens from Black River, however, distinctly incline toward
higginsi. Utterback (1916) has only higginsi from Missouri, but not orhiculata.

It is quite possible that L. higginsi is merely a local form of L. orhiculata,
the form of very large rivers with muddy bottoms. We have repeatedly seen in
other species that inflation of the beaks is a character of big-river-forihs. If this is
another case of this kind, the relation of the Ouachita and Black River orhiculata
to that of the Ohio could be explained as that of an ecological race. Of course,
additional material should be studied. That there might be intergrades is indi-
cated by our specimens from Muscatine, which incline somewhat toward higginsi

As far as I can see L. higginsi (Lea) differs from L. orhiculata chiefly in the more inflated and
more elevated beaks, and in the color of the epidermis, wliich has more green in it.

OETMANN; MONOGEAPH of the naiades of PENNSYLVANIA.

325

by their slightly more elevated beaks and their more brownish-olive epidermis,
and by specimens of the same character from Black River, as well as by a specimen
from the Mississippi at Andulusia, received together with typical orhiculata.

Genus Teuncilla Rafinesque (1820).

Ortmann, 1912, p. 354; Simpson, 1914, p. 2.

Type Truncilla triquetra Rafinesque.

Simpson (1900) has divided the genus into four subgenera {Truncilla, Scale-
naria, Dysnomia, Pilea) while Walker (1910) in a recent revision of the genus,
admits three main groups, which properly might be regarded as subgenera. Investi-
gations of the soft parts, however, probably will require some alterations of this
arrangement. I prefer to ignore the subgenera for the present, since there are
only two species in Pennsylvania, the first one {triquetra) being a true Truncilla,
while the second {rangiana) would fall under Simpson’s Pilea. The rest of the
species are chiefly found in the Tennessee-Cumberland region.

Key to the Pennsylvanian Species of Truncilla,

Ri. Shell with a very distinct posterior ridge, and distinctly truncate posterior slope. Inflation of female
shell narrow, very convex, restricted to the posterior ridge, not differing from the rest of the shell

in texture, and denticulated on the margin. Rays strongly spotted T. triquetra.

02. Shell with a very indistinct posterior ridge, and the posterior slope not truncate. Inflation of the
female shell very broad and flat, occupying the whole postbasal area, with the nacreous layer poorly
developed, not denticulated on the margin. Rays simple, not spotted T. rangiana.

Truncilla triquetra Rafinesque (1820).

Truncilla triquetra Rafinesque, Simpson, 1914, p. 5.

Plate XXI, figs. 3, 4.

Records from Pennsylvania:

Rhoads, 1899 (Ohio River, Coraopolis, Allegheny Co., and Beaver, Beaver Co.; Beaver River, Wampum,
Lawrence Co.)

Ortmann, 19096, p. 188.

Characters of the shell: Small, moderately thick. Outline subovate to sub-
trapezoidal and subtriangular. Anterior margin rounded. Lower margin gently
curved, but sometimes (chiefly in the female) straight or even slightly emarginate
posteriorly. Upper margin very short, passing in old specimens into the obliquely
descending posterior margin without forming an angle (shape triangular), but in
young specimens there is generally an angle (shape trapezoidal). Posterior margin
forming a rather distinct, but more or less rounded, angle with the lower margin.
This angle is situated very low, only a little elevated above the base-line. Beaks

326

MEMOIRS OF THE CARNEGIE MUSEUM.

moderately swollen, and moderately elevated above the hinge-line, located in
front of the middle of the shell, but not very near the anterior end. Beak-sculpture
rudimentary, consisting of about three faint, indistinct bars, which have a tendency
to fall into two loops. Valves greatly swollen, but rather flat upon the sides, convex
anteriorly, and with a very distinct and strong, but blunt, posterior ridge; no
furrow or depression in front of the latter. Behind the posterior ridge the posterior
slope is sharply truncate and flat, being very little elevated in the middle.

Epidermis yellowish, yellowish green, or light green, always with well developed
dark green to blackish rays, which are distinct, broad and bold, and generally
have the tendency to become dissolved into squarish, triangular, or sagittate spots.
In most cases the rays cover all of the lateral faces of the shell, but they are in-
distinct or entirely absent on the posterior slope. In old specimens the rays
disappear toward the lower margin. Concentric bands of color are indicated
chiefly by a tendency to have the spots of the rays arranged in concentric lines.

Hinge well-developed. Pseudocardinals generally two in each valve, but the
anterior tooth in the right valve is small. The teeth are strong, stumpy, ragged.
Interdentum absent. Laterals of medium length, strong. Beak-cavity moderate.
Dorsal muscle-scars in beak-cavity. Adductor-scars distinct, and anteriorly
strongly impressed, less so posteriorly. Nacre always silvery white.

Sexual differences of the shell very strongly marked. The male has a rather
regular, subtriangular outline, with the posterior ridge moderately elevated, the
greatest diameter of the shell situated slightly in front of the ridge, and with a
moderately broad posterior slope, without sculpture, and with the edge of the
shell entire. In the female the marsupial expansion is located on and restricted
to the posterior ridge. In consequence of this the posterior ridge is greatly ele-
vated, and the greatest diameter of the shell is located upon the posterior ridge,
and in old females the diameter of the shell often exceeds the height of the shell
(a proportion quite unique among Naiades). The posterior slope in the female is
greatly enlarged, and in addition there is a sculpture of radial ribs upon the posterior
ridge and posterior slope, which produces a denticulation on the margin of the
shell in this region. This sculpture may be present, in a rudimentary condition,
in old males. The general outline of the female shell is a little different, the pos-
terior end projecting slightly beyond the line of the lower margin, so that there
is a slight concavity of the lower margin in front of the posterior end. The female
shell does not nearly attain the size of the male shell.

ORTMANN: monograph op the naiades of PENNSYLVANIA.

327

L. H. D.

Size: (Males) 1. Kelly, Cat. No. 61.2983 68 mm. 44 mm. 36 mm.

2. Shenango, Cat. No. 61.4097 60 “ 41 “ 31 “

3. Aladdin, Cat. No. 61.3358 52 “ 34 “ 25 “

4. do. “ “ do. .40 “ 27 “ 20 “

(Females) 5. Kelly, Cat. No. 61.2985 (gravid) 45 “ 25 “ 28 “

6. Aladdin, Cat. No. 61.3358 (gravid) 42 “ 26 “ 31 “

7. Aladdin, Cat. No. 61.3358 (gravid) 38 “ 22 “ 19 “

8. Godfrey, Cat. No. 61.3360 (gravid) 37 “ 22 “ 18 “

The largest individuals recorded here are not surpassed by any specimens
from outside of Pennsylvania.

Soft parts (See Ortmann, 1912, p. 355, fig. 27). Glochidia (See Ortmann,
1911, PL 89, fig. 24).

Breeding season: Gravid females were found on Sept. 5, 1913; Sept. 8, 1914;
Sept. 15, 1913; Sept. 16, 1913; Sept. 17, 1913; Sept. 18, 1908; Sept. 27, 1907;
Sept. 27, 1909; Oct. 5, 1908; Then again on May 20, 1914; May 23, 1911; May
24, 1911.

Glochidia have been observed as early as Sept. 17. On May 20 and 23, dis-
charging females have been found. Thus this species is clearly hradytictic.

• Truncilla triquetra.

■ Truncilla rangiana.

+ Do. (Indian garbage heap).

Remarks: The female of this species is so peculiar in shape and color, and in
possessing the remarkable character of denticulations on the margin of the shell.

328

MEMOIES OF THE CAKNEGIE MUSEUM.

that it cannot be confounded with any other species. The male also is quite
well-defined by its subtriangular shape, strong posterior ridge, and truncate pos-
terior slope, and also its color-pattern. However, to a certain degree the male
resembles Amygdalonaias truncata (Rafinesque) but the latter is a higher and
more compressed shell, with the posterior ridge sharper, chiefly in the region of the
beaks, and with the posterior slope narrower and less distinctly truncate.

There is a good deal of variation in the shell of Truncilla triquetra. Aside
from the general outline, which is more subtrapezoidal in young shells, and more
subtriangular in old ones, there is variation in the elongation of the shell. In old
males the posterior end is often considerably drawn out. Both males and females
vary greatly in diameter and in the width of the posterior slope, which in some
old females becomes really freakish. There is also great variation in the color-
pattern, and the rays and the spots are hardly ever alike in any two specimens.

The Lake Erie form of this species does not seem to differ from the normal
type. I have only few specimens from the lake, which compared with specimens
from the Ohio are of about medium size, and seem to be slightly lighter in color.

Localities in Pennsylvania represented in the Carnegie Museum:

Ohio River, Neville Island, Allegheny

Beaver River, Wampum, Lawrence Co. (G. H. Clapp & H. H. Smith).®^^

Mahoning River, Mahoningtown and Edinburg, Lawrence Co.

Shenango River, Pulaski, Lawrence Co.; Sharpsville, Clarksville, and Shenango, Mercer Co.
P3"matuning Creek, Pymatuning Township, Mercer Co.

Allegheny River, Aladdin, Godfrey, Johnetta, Kelly, Mosgrove, and Templeton, Armstrong Co.

Crooked Creek, Rosston, Armstrong Co.

French Creek, Utica, Venango Co.; Meadville and Cambridge Springs, Crawford Co.

Conneaut Outlet, Conneautlake, Crawford Co.

Leboeuf Creek, Waterford, Erie Co.

Dunkard Creek, Mount Morris, Greene Co.

Lake Erie, Presque Isle Bay, Erie, Erie Co.

Other localities represented in the Carnegie Museum:

Lake-drainage:

Sandusky River, Fremont, Sandusky Co., Ohio (C. Goodrich).

Swan Creek, Toledo, Lucas Co., Ohio (C. Goodrich).

Ohio-drainage:

Ohio River, Portland, Meigs Co., Ohio.

Tuscarawas River, Ohio (Holland collection).

Miami River, Ohio (Juny collection).

A dead shell has been seen at Industry, Beaver Co.

A dead shell has been seen in Slipperyrock Creek, at Wurtemberg, Lawrence Co.

ORTMANN: monograph of the naiades of PENNSYLVANIA.

329

Big Beaver Creek, Mercer Co., Ohio (C. Goodrich) (Wabash-drainage).

West Fork River, Lynch Mines, Harrison Co.; West Milford, Harrison Co. (W. F. Graham); Lightburn,
Lewis Co., West Virginia.

Little Kanawha River, Grantsville, Calhoun Co. (W. F. Graham); Burnsville, Braxton Co., West Virginia.
North Fork Hughes River, Cornwallis, Ritchie Co.,. West Virginia.

Tennessee-drainage :

Duck River, Columbia, Maury Co., Tennessee (G. H. Clapp, donor).

Tennessee River, Florence, Lauderdale Co., Alabama (H. H. Smith).

Bear Creek, Burleson, Franklin Co., Alabama (H. H. Smith).

Flint River, Maysville, Madison Co., Alabama (H. H. Smith).

Paint Rock River, Paint Rock, Jackson Co., Alabama (H. H. Smith).

Nolichucky River, Chunns Shoals, Hamblen Co., Tennessee.

Holston River, McMillan and Mascot, Knox Co.; Turley Mill and Holston Station, Grainger Co.;

Austin Mill, Hawkins Co., Tennessee.

South Fork Holston River, Pactolus, Sullivan Co., Tennessee.

North Fork Holston River, Rotherwood, Hawkins Co., Tennessee; Hilton, Scott Co., Virginia; Mendota,
Washington Co., Virginia.

Clinch River, Solway, Knox Co.; Edgemoor and Clinton, Anderson Co.; Black Fox Ford, Union Co.;
Clinch River Station, Claiborne Co.; Oakman, Grainger Co., Tennessee; Speers Ferry and Clinch-
port, Scott Co., Virginia.

Powell River, Combs, Claiborne Co., Tennessee.

Distribution and Ecology in Pennsylvania (See fig. 34) ; This is a widely dis-
tributed species in the Ohio-drainage in western Pennsylvania, but is not found
anywhere in great numbers. In the Ohio proper below Pittsburgh I found only a
few dead shells. It is present in the Beaver-drainage, but is not very abundant.
In the Allegheny in Armstrong Co. it is not rare, and in this region enters some
of the tributaries, as Crooked Creek, and farther up, French Creek, penetrating
well into the headwaters. It has not been found in the Allegheny above Oil City.

In the Monongahela-drainage, this species is extremely rare, and I found it
only in Dunkard Creek. However, farther up in the headwaters in West Virginia
it is rather abuhdant, as in West Fork River, where at Lynch Mines I secured the
largest number of specimens in all my collecting.

In addition this species is present in Lake Erie, but here it is extremely rare.
I found only a single dead shell in Presque Isle Bay.

Truncilla triquetra distinctly belongs to those species, which inhabit riffles.
Wherever I found it, it was on riffles, and (at low stage) in quite shallow water,
in strong currents. Here it is deeply buried, and only the truncated posterior
slope is exposed, so that in the natural position the shell offers a very peculiar
aspect; only the flat, broadly lanceolate posterior slope is exposed to view, differ-
ing entirely from the dark slit generally seen in other Naiades (formed by the

330

MEMOIRS OF THE CARNEGIE MUSEUM.

anal and branchial openings). The bottom at these localities generally consists
of finer or coarser, closely packed gravel; but I have also found this species in pure
sand, when the latter is not kept moving by the current. In Lake Erie, in Presque
Isle Bay, of course, this species must live in the fine sand of the bay : but I did not
find it alive there.

General distribution: Type locality, Falls of the Ohio (Rafinesque) (at Louis-
ville, Jefferson Co., Kentucky).

Simpson (1900) gives the following range: “Ohio River-drainage; western
New York to southern Michigan; Iowa; eastern Nebraska to Indian Territory.”
In the Ohio-drainage, this species is generally distributed, not only in the Ohio
proper, where it seems to be scarce, but also, and chiefly so, in the tributaries.
It goes rather far up in the smaller streams. From Pennsylvania it goes through
West Virginia and Ohio to Indiana and Illinois. Here it goes up into the Kankakee
River and to Will Co. (Baker, 1906), but is not in the “Chicago area.”

Records from the southern tributaries of the Ohio in Kentucky are scarce,
but it is in the upper Cumberland and its tributaries (Wilson & Clark, 1914),
and in the Tennessee to northern Alabama and eastern Tennessee, where it reaches,
in the Clinch and North Fork of the Holston River, into the state of Virginia.

Farther to the west, it has been reported from the Mississippi River in Iowa
(Pratt, 1876; Witter, 1878; Marshall, 1895), but not farther north. Simpson
records it from eastern Nebraska, and it exists in eastern Kansas (Scammon, 1906),
and also, according to Simpson, in Oklahoma. However, there are no records
from Missouri (see XJtterback, 1916) and Arkansas, and southward, and none from
the Alabama-drainage.

T. triquetra has, however, crossed over into the lake-drainage, and it is found
in this drainage in Ohio (Sterki, 1907a), in southern Michigan (Walker, 1898),
and in Lake Erie (Walker, 1913). Here it goes eastward to the state of New York,
and the locality “New York” (Call, 1885; Simpson, 1900) is supported only by
definite records from Buffalo and Niagara River (see Marshall, 1895).^^^

As in Pennsylvania so elsewhere this species seems to prefer riffles, with rough
bottoms and strong currents.

It might be in the upper Allegheny in New York, but, as has been said, I never found it in the
Allegheny above Oil City.

OETMANN: monograph of the naiades of PENNSYLVANIA.

331

Truncilla rangiana (Lea) (1838).

Truncilla perplexa rangiana (Lea) Simpson, 1914, p. 25.^^®

Plate XXI, figs. 5, 6, 7.

Records from Pennsylvania:

Marshall, 1896 (as U. Lea from the Allegheny River, Warren Co.; this undoubtedly is T.

rangiana).

Ortmann, 19096, p. 188.

Characters of the shell: Shell rather small (but larger than that of T. triquetra),
moderately thick, but extremely thin at the postbasal expansion of the female.
Outline irregularly subovate. Anterior margin rounded. Lower margin of the
male convex in front, straight or concave, and ascending in the posterior part;
this posterior part forms a blunt, more or less distinct angle with the anterior part.
In the female the lower margin forms a more or less distinctly double curve, the
posterior part more strongly convex, and the two parts are separated by a short
concavity or notch. Upper margin moderate, passing in a blunt angle or gradually
into the obliquely descending posterior margin. In the male the posterior and
lower margins meet in a blunt posterior point, which is situated well above the
basal line; in the female the posterior end of the shell is broadly and evenly rounded.
Beaks not much swollen, and little elevated above the hinge-line, located in front
of the middle of the shell, and only a short distance away from the anterior end.
Beak-sculpture rudimentary, consisting of three or four faint bars, which are
double-looped. Valves rather compressed, moderately and rather evenly convex
in the anterior part of the shell. In the male there is a blunt and indistinct pos-
terior ridge, and in front of this a broad and shallow radial furrow, running from
the beak to the posterior part of the lower margin. In front of this furrow the
shell is often somewhat elevated, forming a blunt rib running toward the angle
in the middle of the lower margin. Posterior slope narrow, gently convex or
flattened, not truncate. In the female the region of the furrow and of the posterior
ridge is occupied by a broad and somewhat flattened expansion, in front of which
there is a more or less distinct, narrow constriction in old specimens. Posterior
ridge and posterior slope entirely indistinguishable in the female.

Epidermis yellow or greenish olive, generally with rather distinct, fine, and
crowded, straight and uninterrupted, dark green rays. These rays may cover
practically the whole surface, but in old specimens they disappear toward the
lower margin and upon the posterior expansion, which often is of a lighter color

U. gubernaculum Reeve, 1865, PL 28, fig. 146, is not this form, as Simpson believes, but a variety
of T. torulosa (Rafinesque) (= perplexa Lea) found in the headwaters of the Tennessee River.

332

MEMOIRS OF THE CARNEGIE MUSEUM.

than the rest of the surface. Concentric dark bands, marking growth-rests, are
more or less distinct.

Hinge well-developed. Pseudocardinals two in left, one or two in right valve,
stumpy, ragged. Interdentum absent. Laterals moderately long, moderately
strong. Beak-cavity rather shallow. Dorsal muscle-scars in beak-cavity. Ad-
ductor-scars distinct and well impressed, chiefly those in front. Nacre always
white.

Sexual differences in the shell extremely great. The general form of the male
shell is ovate, somewhat pointed posteriorly, with a rather distinct posterior ridge
and a radial furrow in front of it. The female shell is also ovate, but the wider
part is behind, and it is not at all pointed at the posterior end, but broadly rounded.
The place of the radial furrow is filled by a broad postbasal expansion, which may
occupy the whole posterior half of the shell, and from the anterior part, this is
marked off, in old specimens, by a more or less distinct constriction. In young
females (PI. XXI, fig. 6), this postbasal expansion is less developed, and the shell
is only slightly thinner here than the rest of it. But in old females this expansion
is greatly developed, and is very thin, since the deposition of nacreous matter
ceases here, and thus the expansion appears chiefly horny (epidermis), with a thin
film of prismatic matter. The females gi’ow practically to the same size as the

males.

L. H. D.

Size: (Males) I. Meadville, Cat. No. 61.3362 72 mm. 50 mm. 31 mm.

2. Cochranton, Cat. No. 61.3363 70 “ 50 “ 33 “

3. do. “ “ do 61 “ 46 “ 29 “

4. do. “ “ do. . 45 “ 33 “ 22 “

5. Utica, Cat. No. 61.3368 37 “ 28 “ 19 “

(Females) 6. Warren, Cat. No. 61.3979 70 “ 45 “ 32 “

7. Cochranton, Cat. No. 61.3363 (gravid) 65 “ 43 “ 30 “

8. Hickory, Cat. No. 61.3366 (gravid) . .62 “ 42 “ 25 “

9. Cochranton, Cat. No. 61.3363 ....51 “ 36 “ 25 “

10. Harbor Bridge, Cat. No. 61.3372 . .36 “ 26 “ 17 “

Soft 'parts (See Ortmann, 1912, p. 358, fig. 28). Glochidia (See ibid.) similar
to those of T. perplexa Lea (Obs. VI, 1858, PL 5, fig. 21) (= torulosa Rafinesque).

Breeding season: Gravid females were collected on Sept. 2, 1908; Sept. 5,
1908; Sept. 6, 1908; Sept. 15, 1909; Sept. 18, 1908. On the first date young
glochidia were already observed, and thus the beginning of the season probably
falls early in August. The species probably is hradytictic.

Remarks: T. rangiana has been considered a variety of T. torulosa (Rafin-
esque) = perplexa Lea. The latter is not found in Pennsylvania. It is dis-

ORTMANN: monograph of the naiades op PENNSYLVANIA.

333

tinguished from T. rangiana by the presence of a radial row of tubercles upon the
middle of the shell, standing in the male upon the radial rib, in the female in front
of the marsupial expansion; and further T. torulosa has the postbasal expansion
of the female dark green in color.

Slightly tuberculate specimens once existed in Pennsylvania, and I collected
such in an Indian Garbage heap at Point Marion (Ortmann, 1909c). I have
called them T. cincinnatiensis, but they are not the true cincinnatiensis, and are
by no means the typical T. torulosa. Whether forms approaching the latter once
existed in the Monongahela and Ohio cannot be now ascertained. All living
shells from Pennsylvania are typical rangiana. Some specimens, indeed, show
indications of tubercles, but these are rather indefinite, obscure swellings, the
majority of the specimens being perfectly smooth.

The female of this species is easily recognized by the peculiar postbasal ex-
pansion, The male, however, has a shape which reminds of young Plethobasus
cyphyus. But P. cyphyus has a lighter yellow epidermis, and has no rays. More-
over the beak-sculpture is quite different.

The range of variation has-been already indicated in the description. Dis-
regarding the slight tendency to show traces of tubercles in the middle of the
shell, the radial furrow is somewhat variable, and also the angle in the middle
of the lower margin. The postbasal expansion of the female is very variable,
increasing with the age of the individual, but it always is of light color, not dark-
green.

Localities in Pennsylvania represented in the Carnegie Museum:

Shenango River, Harbor Bridge and Pulaski, Lawrence Co.

Allegheny River, Aladdin, Godfrey, Johnetta, and Templeton, Armstrong Co.; Walnut Bend, Venango
Co.; Tionesta and Hickory, Forest Co.; Warren, Warren Co.

French Creek, Utica, Venango Co.; Cochranton and Meadville, Crawford Co.

Connewango Creek, Bussell, Warren Co.

Other localities represented in the Carnegie Museum:

West Fork River, Lynch Mines, Harrison Co., West Virginia.

Tuscarawas River, Ohio (Holland collection).

Columbus, Franklin Co., Ohio (Hartman collection) (Smith collection).

White River, Rockford, Jackson Co., Indiana (G. H. Clapp, donor) (Wabash-drainage).

Distribution and Ecology in Pennsylvania (See fig. 34); I have found this
species in the Beaver-drainage in the Shenango River,^^®“ where it is very Fare

I have not seen it in the Mahoning River in Pennsylvania, although it has been reported from
this river in Ohio (Lea, 1838; Dean, 1890).

334

MEMOIES OF THE CAENEGIE MUSEUM.

(only three specimens taken), and in the Allegheny River from Armstrong County
up to Warren County. It is also found in French and Connewango Creeks,
tributaries of the Allegheny. In the Allegheny itself it is found rather regularly,
but not in great numbers; it is most abundant in French Creek and at Cochranton
it was a common species.

In the Monongahela-drainage I never found it alive, but its former occurrence
there is indicated by shells from the Indian garbage heap at Point Marion. It
also occurs farther up in the headwaters in West Virginia, but is very rare.

It is remarkable that this species has not turned up in the Ohio below Pitts-
burgh, and that there are no localities for it' positively ascertained in the Ohio,
until we reach Cincinnati.

I always found this species on riffles, perferably upon a bottom composed of
firmly packed and rather fine gravel, in swiftly flowing, shallow water. In the
Allegheny in Armstrong County I have also seen it in coarse gravel.

General distribution: Type locality, Ohio River, Cincinnati (Lea).

Aside from western Pennsylvania and the upper Monongahela in West Vir-
ginia, this species is known from the state of Ohio. In the Ohio proper only the
type locality is known; but according to Sterki (1907a) it is found in the Mahoning,
Tuscarawas, and Scioto Rivers. In Indiana it is found in the Ohio and Wabash
(Call, 1896a), and in White River (Carnegie Museum) and in Illinois only in the
southern portion (Wabash, see Baker, 1906). In addition it occurs in_ south-
eastern Michigan, in the Lake Erie-drainage (Detroit and Raisin Rivers, see Walker,
1892 & 1898), and it is also given (Walker, 1913) in the Lake Erie list, but has
never been found on the Pennsylvanian shores of the lake.

According to these records, this species is rather restricted in its range, and
belongs to the Ohio River and its tributaries, chiefly the northern ones, and seems
to have its metropolis in the smaller tributaries of the upper Ohio River in the
states of Ohio and Pennsylvania. It also has crossed over into the lake-drainage,
but particulars are lacking.

An allied species is T. tondosa (= perplexa), which seems to inhabit the
lower Ohio and the Tennessee drainage. It is interesting to note that the latter
develops in the upper Tennessee a compressed form without tubercles, which should
be called gubernaculum Reeve, and which is a form parallel to rangiana, but has
the dark green postbasal expansion of torulosa. This case should be kept in mind,
for it is an interesting instance of the reactions of the Naiades to environmental
conditions. The form most closely related to T. rangiana undoubtedly is T.
sampsoni (Lea) of the lower Wabash, which differs only in having the shell more

ORTMANN: monograph of the naiades of PENNSYLVANIA.

335

swollen and the beaks more inflated. It possibly is only the big-river-form of

T. rangiana.

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340

MEMOIES OF THE CAENEGIE MUSEUM.

1909a Ortmann, A. E. The Breeding Season of Unionidse in Pennsylvania (Nautilus,
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19096 Ortmann, A. E. A preliminary List of the Unionidse of Western Pennsylvania,
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1909c Ortmann, A. E. Unionidse from an Indian Garbage Heap (Nautilus, XXIII,
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1910 Ortmann, A. E. A New System of the Unionidse (Nautilus, XXIII, 1910, pp.
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1911a Ortmann, A. E. The Anatomical Structure of Certain Exotic Najades compared
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19116 Ortmann, A. E. Monograph of the Najades of Pennsylvania (Memoirs Car-
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1912 Ortmann, A. E. Notes upon the Families and Genera of the Najades (Annals
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1913a Ortmann, A. E. The Alleghenian Divide, and its Influence upon the Fresh-
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19136 Ortmann, A. E. Studies in Najades (Nautilus, XXVII, 1913, pp. 88-91).

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1901 PiLSBRY, H. A. Alasmidonta marginata (Say) (Nautilus, XV, 1901, pp. 16-17).

1876 Pratt, W. H. List of Shells at Davenport, Iowa (Proceedings Davenport Academy
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1820 Pafinesque, C. S. Monographie des Coquilles Bivalves et Fluviatiles de la
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1899 Rhoads, S. N. On a Recent Collection of Pennsylvania Mollusks from the Ohio
River System below Pittsburgh (Nautilus, XII, 1899, pp. 133-137).

1904 Rhoads, S. N. A Glimpse at the Shell Fauna of Delaware (Nautilus, XVIII,
1904, pp. 63-67).

Richardson, R. E. (See Forbes & Richardson.)

ORTMANN: monograph of the naiades of PENNSYLVANIA.

341

190* Ruthven, a. G. Notes on the Molluscs, Reptiles, and Amphibians of Ontonagon
County, Michigan {Sixth Annual Report of the Michigan Academy of Science,
p. 191).

Say, T. (See Binney, W. G.)

1906 Scammon, R. E. The Unionidse of Kansas {Kansas University Bulletins, III,
1906, pp. 279-373).

1912 Scharff, R. F. Distribution and Origin of Life in America. New York. 1912.
1895 Schick, M. Mollusk Fauna of Philadelphia and Environs {Nautilus, VIII, 1895,
pp. 133-140).

1891 Simpson, C. T. Notes on Unionidse {Nautilus, V, 1891, pp. 86-88).

1893 Simpson, C. T. On Some Fossil Unios and Other Shells from the Drift at Toronto
{Proceedings U. S. National Museum, XVI, 1893, pp. 591-595).

1895 Simpson, C. T. Unio ochraceus and cariosus {Nautilus, VIII, 1895, pp. 121-
123).

1900 Simpson, C. T. Synopsis of the Naiades, or Pearly Fresh-water Mussels {Pro-
ceedings U. S. National Museum, XXII, 1900, pp. 501-1044).

1914 Simpson, C. T. A Descriptive Catalogue of the Naiades, or Pearly Fresh-water

Mussels, Detroit, 1914.

Simpson, C. T. (See Dale & Simpson.)

1893 SiNGLEY, J. A. Texas Molluscs {Fourth Annual Report of the Geological Survey

of Texas for 1892 (1893), p. 316).

1899 Smith, H. M. The Mussel Fishery and Pearl-button Industry of the Mississippi
River {Bulletin U. S. Fish Commission, XVIII, for 1898 (publ. 1899-), pp. 289-
314).

SowERBY, G. B. (See Reeve, L. A.)

1898 Sterki, V. Some Observations on the Genital Organs of the Unionidse, with
Reference to Classification {Nautilus, XII, 1898, pp. 18-32).

1907a Sterki, V. A Preliminary Catalogue of the Land and Fresh-water Mollusca
of Ohio {Proceedings of the Ohio State Academy of Sciences, IV, 1907, pp. 367-
402).

19075 Sterki, V. Note on Tritogonia tuberculata {Nautilus, XXI, 1907, p. 48).

1894 Stupakoff, S. H. Land and Fresh-water Shells of Allegheny County, Pennsyl-

vania {Nautilus, VII, 1894, pp. 135-136).

1912 SuRBER, T. Identification of the Glochidia of Fresh- water Mussels {U. S. Bureau
of Fisheries, Doc. 771, 1912, pp. 1-10).

1915 SuRBER, T. Identification of the Glochidia of Fresh-water Mussels {U. S. Bureau

of Fisheries, Doc. 813, 1915, pp. 1-9).

SuRBER, T. (See Coker & Surber.)

* Separate copy, without date, but published after 1903.

MEMOIRS OF THE CARNEGIE MUSEUM.

342

1868 Tryon, G. W. Notes on Mollusca Collected by F. V. Hayden in Nebraska
{American Journal Conchology, VI, 1868, pp. 150-151).

1916 Utterback, W. I. The Naiades of Missouri {American Midland Naturalist,
IV, 1916, pp. 1-200).

1910 Vanatta, E. G. Unionidse from Southeastern Arkansas and N. E. Louisiana
{Nautilus, XXIII, 1910, pp. 102-104).

1915 . Vanatta, E. G. Rafinesque’s Types of Unio {Proceedings Academy of Natural
Sciences of Philadelphia, 1915, pp. 549-559).

1893 Vaughan, T. W. Notes on Mollusks from Northwestern Louisiana and Harrison

County, Texas {American Naturalist, XXVIII, 1893, pp. 944-961).

1892 Walker, B. The Shell-bearing Mollusca of Vlichigan {Nautilus, VI, 1892, p.
42 et seq.).

1894 Walker, B. Shells of Saginaw Valley, Michigan {Nautilus, VII, 1894, pp. 125-

129).

1898 Walker, B. The Distribution of the Unionidse in Michigan {Report of Michigan
Academy of Science, 1898, pp. 1-20).

1909 Walker, B. Annotated List of the Mollusca of Isle Royale, Michigan (In
Adams, C. C., An Ecological Survey of Isle Royale, Lake Superior {Report
University of Michigan Museum, 1909, pp. 281-298).

1910a Walker, B. The Distribution of Margaritana margaritifera (L.) in North
America {Proceedings Malacological Society of London, IX, 1910, pp. 126-145).
19106 Walker, B. On the Validity of Unio undatus Barnes {Nautilus, XXIV, 1910,
pp. 6-10; 16-24).

1910c Walker, B. Notes on Truncilla, with a Key to the Species {Nautilus, XXIV,
1910, pp. 75-81). I

1913 Walker, B. The Unione Fauna of the Great Lakes {Nautilus, XXVII, 1913,
pp. 18-23; 29-34; 41-47; 56-59).

1915 Walker, B. Unio viridis Conrad. {Nautilus, XXIX, 1915, pp. 74-78.)

1916 Walker, B. The Rafinesque-Poulson Unios {Nautilus, XXX, 1916, pp. 43-47).
1918 Walker, B. Notes on North American Naiades. I. {Occasional Papers of

the Museum of Zoology of the University of Michigan, No. 49, 1918, pp. 1-6.)
1908 Walker, R. D. & Coolidge, W. H., Jr. Mollusca of Keene, New Hampshire
{Nautilus, XXII, 1908, p. 32).

1918 Wheeler, H. E. The Mollusca of Clark Co., Arkansas {Nautilus, XXXI, 1918,
pp. 109-125).

1863 Whiteaves, J. F. On the Land and Fresh-water Mollusks of Lower Canada
{Canadian Naturalist and Geologist, VIII, 1863, p. 99 et seq.)

1912a Wilson, C. B. & Clark, H. W. The Mussel Fauna of the Kankakee Basin
{U. S. Bureau of Fisheries, Doc. 758, 1912, pp. 1-52).

ORTMANN: monograph of the naiades of PENNSYLVANIA.

343

19126 Wilson, C. B. & Clark, H. W. Mussel Beds of the Cumberland River in 1911
(U. S. Bureau of Fisheries, Economic Circular, I, 1912, pp, 1-4).

1914 Wilson, C. B. & Clark, H. W. The Mussels of the Cumberland River and its
Tributaries (U. S. Bureau of Fisheries, Doc. 781, 1914, pp. 1-61).

1917 Winslow, M. L. An Annotated List of Shells from Northern Michigan (Occa-
sional Papers of the Museum of Zoology of the University of Michigan, No. 4^,
1917, pp. 1-16).

1878 Witter, F. M. List of the Shells of Iowa (Quarterly Journal of Conchology, I,
1878, pp. 385-394).

344

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE I.

All figures are three-fourths natural size, a, lateral view; b, dorsal view.

Fig. 1. Margaritana margaritif era (Linnseus). Normal specimen from Indian Fun,
Rene Mont, Schuylkill Co., Pennsylvania, collected May 4, 1909; C. AI. Cat. No. 61.4272,

Fig. 2. Fusconaia subrotunda (Lea). Young specimen of typical shape, diameter
64 per cent of length, from Allegheny River, Godfrey, Armstrong Co., Pennsylvania,
collected October 5, 1908; C. M. Cat. No. 61.3941a.

Fig. 3. Fusconaia subrotunda kirtlandiana (Lea). Old female, diameter 45 per
cent of length, slightly drawn out posteriorly, from Allegheny River, Kelly, Armstrong
Co., Pennsylvania, collected August 5, 1913; C. AI. Cat. No. 61.6462.

Fig. 4. Fusconaia subrotunda kirtlandiana (Lea). Typical young specimen, dia-
meter 45 per cent of length. Alay be regarded as a topotype, from Mahoning River,
Mahoningtown, Lawrence Co., Pennsylvania, collected August 4, 1908; C. M. Cat.
No. 61.3921.

Fig. 5. Fusconaia subrotunda kirtlandiana (Lea). Adult specimen representing
the typical phase, diameter 42 per cent of length, also topotype from Mahoning River,
Mahoningtown, Lawrence Co., Pennsylvania, collected October 15, 1908; C. M. Cat.
No. 61.3928.

Memoirs Carnegie Museum, Vol. VIII.

Plate I.

Margaritana-Fusconaia.

/

*1*

ts

. » fv/ --
' I

J

346

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE II.

All figures are three-fourths natural size, a, lateral view; b, dorsal view.

Fig. 1. Fusconaia fiava trigona (Lea). Typical specimen of this form, diameter
66 per cent of length, from Alonongahela River, Charleroi, Washington Co., Pennsyl-
vania, collected by G. A. Ehrmann; C. M. Cat. No. 61.58696.

Fig. 2. Fusconaia flava parvula Grier. Alale, from the type-set, diameter 52 per
cent of length, from Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected
July 8, 1910; C. M. Cat. No. 61.4513.

Fig. 3. Fusconaia flava (Rafinesque). Typical specimen, diameter 42 per cent of
length, from Raccoon Creek, New Sheffield, Beaver Co., Pennsylvania, collected October
11, 1907; C. M. Cat. No. 01.3151.

Fig. 4. A77ihlema plicata (Say). Old full-grown specimen of the typical plicata
from the type-locality. Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, col-
lected June 3, 1908; C. M. Cat. No. 61.3336.

Fig. 5. Amblema plicata (Say). Young specimen, topotype, from Lake Erie,
Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected July 8, 1910; C. M. Cat. No.
61.4518.

Fig. 6. Amblema plicata (Say). Specimen of medium size, unusually compressed,
topotype from Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected
July 8, 1910; C. M. Cat. No. 61.4518.

Fig. 7. Amblema plicata costata (Rafinesque). Rather young specimen of the
much compressed form of the small creeks, from French Creek, Cambridge Springs,
Crawford Co., Pennsylvania, collected September 13, 1909; C. M. Cat. No. 61.4348.

Memoirs Carnegie Museum, Vol. VIII.

Plate II,

Fusconaia-A mblema .

y-::

~.r* • • • ••

T*T

348

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE III.

All figures are three-fourths natural size, a, lateral view; 6, dorsal view.

Fig. 1. Amblema plicata costata (Rafinesque) . Typical, gravid female, of good
size, representing the compressed creek-form, from Pymatuning Creek, Pymatuning
Township, Mercer Co., Pennsylvania, collected July 8, 1909; C. M. Cat. No. 61.4340.

Fig. 2. Amblema plicata costata (Rafinesque). Exceptional specimen, without
oblique undulations, also distinctly more swollen than the normal form, from the Alle-
gheny River, Kelly, Armstrong Co., Pennsylvania, collected August 3, 1908; C. M. Cat.
No. 61.3829.

Fig. 3. Amblema plicata costata (Rafinesque). Young specimen with undulations
nearly obliterated, also somewhat swollen, from Ohio River, Cook’s Ferry, Beaver Co.,
Pennsylvania, collected October 8, 1909; C. M. Cat. No. 61.4428.

Fig. 4. Quadrula pustulosa (Lea). Specimen with few tubercles, from the Mahon-
ing River, Edinburg, Lawrence Co., Pennsylvania, collected September 15, 1908; C. M.
Cat. No. 61.3875.

Fig. 5. Quadrula pustulosa (Lea). Specimen with numerous tubercles, from Ohio
River, Industry, Beaver Co., Pennsylvania, collected October 22, 1908; C. M. Cat.
No. 61.3878.

Fig. 6. Quadrula pustulosa schoocraftensis (Lea). Old female, unusually trans-
verse, from Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected July 8,
1910; C. M. Cat. No. 61.4515

Fig. 7. Quadrula pustulosa schooler aftensis (Lea). Female of medium size and
rather normal shape, from Lake Erie, Cedar Point, Erie Co., Ohio, collected by Chas.
Brookover, August 1909; C. M. Cat. No. 61.4451.

Memoirs Carnegie Museum, Vol, VIII.

Plate III.

Amblema-Quadrula.

350

MEMOIES OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE IV.

All figures are three-fourths natural size, a, lateral .view; 6, dorsal view.

Fig. 1. Quadrula quadrula (Rafinesque) . Only specimen ever found in the Ohio
in Pennsylvania, from Ohio River, Cook’s Ferry, Beaver Co., Pennsylvania, collected
September 10, 1908; C. M. Cat. No. 61.3869.

Fig. 2. Quadrula verrucosa (Rafinesque). Male of medium size, from Shenango
River, Pulaski, Lawrence Co., Pennsylvania, collected July 19, 1909; C. M. Cat. No.
61.4121.

Fig. 3. Quadrula verrucosa (Rafinesque). Rather large female, from Shenango
River, Pulaski, Lawrence Co., Pennsylvania, collected July 19, 1909; C. M. Cat. No.
61.4121.

Fig. 4. Quadrula metanevra (Rafinesque). Normal specimen from Ohio River,
Industry, Beaver Co., Pennsylvania, collected September 25, 1908; C. M. Cat. No.
61.3865.

Fig. 5. Quadrula metanevra (Rafinesque). Specimen intergrading toward the var.
wardi (Lea), with the tubercles moderately developed and the shell slightly more com-
pressed, from Allegheny River, Kelly, Armstrong Co., Pennsylvania, collected September
27, 1907; C. M. Cat. No. 61.3071.

Fig. 6. Quadrula metanevra (Rafinesque). A rather typical wardi (Lea), strongly
compressed and with the large tubercles obhfcerated, from the Allegheny River, Kelly,
Armstrong Co., Pennsylvania, collected August 3, 1908; C. M .Cat. No. 61.3861.

Memoirs Carnegie Museum, Vol. VIII,

Plate IV.

Quadrula.

352

MEMOIES OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE V.

All figures are threte-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Quadrula cylindrica (Sa-y). Normal individual, from the Allegheny River,
Godfrey, Armstrong Co., Pennsylvania, collected August 10, 1909; C. M. Cat. No.
61.4358.

Fig. 2. Quadrula cylindrica (Say), Intergrade toward the smooth form, with
tubercles greatly reduced, but shell not much compressed, from French Creek, Coch-
ranton, Crawford Co., Pennsylvania, collected September 2, 1908; C. M. Cat. No.
61.3851.

Fig. 3. Quadrula cylindrica (Say). Smooth and compressed form, characteristic
for the headwaters of the Ohio, from French Creek, Meadville, Crawford Co., Penn-
sylvania, collected September 1, 1908; C. M. Cat. No. 61.3850.

Fig. 4. Rotundaria tuberculata (Rafinesque) . Female from the Allegheny River,
Godfrey, Armstrong Co., Pennsylvania, collected August 10, 1909; C.' M. Cat. No.
61.4411.

Fig. 5. Plethohasus cooperianus (Lea). Normal individual from the Ohio River,
Industry, Beaver Co., Pennsylvania, collected October 22, 1908; C. M. Cat. No. 61.3882.

Fig. 6. Plethohasus cyphyus (Rafinesque). Large specimen of normal shape,
from the Allegheny River, Kelly, Armstrong Co., Pennsylvania, collected September 27,
1907; C. M. Cat. No. 61.3065.

Memoirs Carnegie Museum, Vol. VIII.

Plate V.

■ t..

Quadrula-Rotundaria-Plethobasus.

354

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE VI.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Plethobasus cyphyus (Rafinesque) . Same specimen as the one figured on
Plate V, fig. 6, from the Allegheny River, Kelly Armstrong Co., Pennsylvania, collected
September 27, 1907; C. M. Cat. No. 61.3065.

Fig. 2. Plethobasus cyphyus (Rafinesque). Specimen of medium size, and rather
short, from the Allegheny River, Kelly, Armstrong Co., Pennsylvania, collected Sep-
tember 27, 1907; C. M. Cat. No. 61.3065.

Fig. 3. Plethobasus cyphyus (Rafinesque). Small specimen from the Allegheny
River, Kelly, Armstrong Co., Pennsylvania, collected September 27, 1907; C. M. Cat.
No. 61.3065.

Fig. 4. Pleurobema obliquum (Lamarck). Normal form of the upper Ohio,
diameter 55 percent of length, from the Ohio River, Industry, Beaver Co., Pennsylvania,
collected October 3, 1908; C. M. Cat. No. 61.3900.

Fig. 5. Pleurobema obliquum (Lamarck). Specimen with weak furrow, inclining
toward the var. catillus (Conrad), diameter 57 percent of length, from the Ohio River,
Industry, Beaver Co., Pennsylvania, collected September 25, 1908; C. M. Cat. No.
61.3894.

Fig. 6. Pleurobema obliquum catillus (Conrad). A specimen closer to catillus
than Fig. 5, diameter 56 percent of length, nacre pink. Specimens like those given in
figs. 5 and 6 demonstrate that the two forms intergrade. From the Ohio River, In-
dustry, Beaver Co., Pennsylvania, collected September 8, 1908; C. M. 'Cat. No. 61.3895.

Fig. 7. Pleurobema obliquum rubrum (Rafinesque). Rather typical, diameter 57
percent of length, from the Allegheny River, Godfrey, Armstrong Co., Pennsylvania,
collected July 13, 1908; C. M. Cat. No. 61.3891.

Fig. 8. Pleurobema obliquum (Lamarck). Old specimen, drawn out posteriorly
and consequently with the low diameter of 42 percent of the length; when young,
this was a rather tj^pical obliquum. From the Ohio River, Industry, Beaver Co., Penn-
sylvania, collected October 3, 1908; C. M. Cat. No. 61.3900.

Memoirs Carnegie Museum, Vol, VIII.

Plate VI,

Plethobasus-Pleurohema.

356

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE VII.

All figures are three-fourths natural size, a, lateral view; b, dorsal view.

Fig. 1. Pleurobema obliquum cordatum (Rafinesque) . This specimen most closely
approaches this form, but is not quite typical. From the Allegheny River, Godfrey,
Armstrong Co., Pennsylvania, collected August 10, 1909; C. M. Cat. No. 61.4378.

Fig. 2. Pleurobema obliquum catillus (Conrad). Female, diameter 52 percent of
length, approaching the var. coccineum (Conrad). From the Allegheny River, Godfrey,
Armstrong Co., Pennsylvania, collected July 27, 1910; C. AI. Cat. No. 61.4567.

Fig. 3. Pleurobema obliquum coccineum (Conrad). Large specimen, diameter 42
percent of length, from French Creek, Cambridge Springs, Crawford Co., Pennsylvania,
collected September 13, 1909; C. AI. Cat. No. 61.4394.

Fig. 4. Pleurobema obliquum coccineum (Conrad). Younger specimen, diameter
45 percent of length, from French Creek, Cambridge Springs, Crawford Co., Pennsyl-
vania, collected September 13, 1909; C. AI. Cat. No. 61.4394.

Fig. 5. Pleurobema obliquum coccineum (Conrad). Topotype, diameter 46 per-
cent of length, from Alahoning River, Alahoningtown, Lawrence Co., Pennsylvania,
collected August 4, 1908; C. AI. Cat. No. 61.3907.

Fig. 6. Pleurobema obliquum pauperculum (Simpson). Typical, large specimen,
diameter 47 percent of length, from Lake Erie, Presque Isle Bay, Erie, Erie Co., Penn-
sylvania, collected Alay 21, 1909; C. AI. Cat. No. 61.4398.

Fig. 7. Pleurobema clava (Lamarck). Large, gravid female from Neshannock
Creek, Eastbrook, Lawrence Co., Pennsylvania, collected June 18, 1908; C. AI. Cat. No.
61.3335.

Fig. 8. Pleurobema clava (Lamarck). Smaller specimen, very oblique, from
Shenango River, Clarksville, Afercer Co., Pennsylvania, collected September 21, 1908;
C. AI. Cat. No. 61.3814.

Fig. 9. Pleurobema clava (Lamarck). Small specimen, much less oblique, from
Shenango River, Clarksville, Alercer Co., Pennsylvania, collected September 21, 1908;
C. AI. Cat. No. 61.3814.

Note: The rays and blotches present in the specimens in figs. 7, 8, and 9, did not
come out in the photographs.

Memoirs Carnegie Museum, Vol. VIII.

Plate VII.

Pleurobema.

358

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE VIII.

All figures are three-fourths natural size, a, lateral view; 6, dorsal view.

Fig. 1. Elliptio niger (Rafinesque) . Normal specimen of medium size, from the
Ohio River, Industry, Beaver Co., Pennsylvania, collected September 8, 1908; C. M.
Cat. No. 61.3781.

Fig. 2. Elliptio dilatatus (Rafinesque) . Gravid female of medium size, diameter
29 percent of length, from the Allegheny River, Godfrey, Armstrong Co., Pennsylvania,
collected July 27, 1910; C. M. Cat. No. 61.4618.

Fig. 3. Elliptio dilatatus sterkii Grier. Female, one of the types, which, however,
is not much swollen, the diameter being only 32 percent of the length. From Lake
Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected July 12, 1910; C. M.
Cat. No. 61.4628.

Fig. 4. Elliptio violaceus (Spengler). Normal specimen as found in smaller
streams, from a small tributary of Lizard Creek, West Penn, Schuylkill Co., Pennsylvania,
collected May 4, 1909; C. M. Cat. No. 61.4308.

Fig. 5. Elliptio violaceus (Spengler). Strongly compressed form with rough
epidermis corresponding to the form called jejunus by Lea, from a mill-race of Crooked
Creek, Tioga, Tioga Co., Pennsylvania, collected August 19, 1909; C. M. Cat. No.
61.4312.

Fig. 6. Elliptio cupreus (Rafinesque). Specimen of medium size from Great
Tonoloway Creek, Thompson Township, Fulton Co., Pennsylvania, collected September
5, 1909; C. M. Cat. No. 61.4322.

Fig. 7. Elliptio fisherianus (Lea). Normal specimen of medium size from Chop-
tank Mills, Kent Co., Delaware, collected by S. N. Rhoads, June, 1903; C. M. Cat. No.
61.4645.

Memoirs Carnegie Museum, Vol. VIII.

Plate VIII.

ElUptio.

360

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE IX.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Lasmigona (Platynaias) viridis (Rafinesque). Gravid specimen, about

full-grown, from Conneaut Outlet, Conneautlake, Crawford Co., Pennsylvania, collected
May 14, 1908; C. M. Cat. No. 61.3297.

Fig. 2. Lasmigona (Patynaias) viridis (Rafinesque). Gravid specimen, young.
This specimen shows the great similarity of young L. viridis to L. subviridis (compare
Fig. 4). From Conneaut Outlet, Conneautlake, Crawford Co., Pennsylvania, collected
August 7, 1908; C. M. Cat. No. 61.3679.

Fig. 3. Lasmigona {Platynaias) subviridis (Conrad). Gravid specimen, about

full-grown, with exceptionally solid shell, from water heavily charged with lime. From
Conococheague Creek, Greencastle, Franklin Co., Pennsylvania, collected September 6,
1909; C. M. Cat. No. 61.4222.

Fig. 4. Lasmigona {Platynaias) subviridis (Conrad). Gravid specimen of the

normal, thin-shelled form, from Schuylkill Canal, Manayunk, Philadelphia Co., Penn-
sylvania, collected April 24, 1909; C. M. Cat. No. 61.4219.

Fig. 5. Lasmigona {Lasmigona) costata (Rafinesque). Gravid female of typical

shape, from Shenango River, Shenango, Mercer Co., Pennsylvania, collected September
27, 1909; C. M. Cat. No. 61.4235.

Fig. 6. Lasmigona {Lasmigona) costata eriganensis Grier. Specimen from the
set of types. From Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected
May 22, 1909; C. M. Cat. No. 61.4223.

Fig. 7. Lasmigona {Pterosyna) complanata (Barnes). Gravid female, not quite
full-grown, from Leboeuf Creek, Waterford, Erie Co., Pennsylvania, collected September
14, 1909; C. M. Cat. No. 61.4238.

Memoirs Carnegie Museum, Vol VIII.

Plate IX.

Lasmigona.

362

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE X.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Anodonta grandis Say. Normal creek-form, from Crooked Creek, Creek-
side, Indiana Co., Pennsylvania, collected May 27, 1908; C. M. Cat. No. 61.3649.

Fig. 2. Anodonta grandis Say. Gravid female of the pond-form gigantea (Lea),
from pond at Harmarville, Allegheny Co., Pennsylvania, collected October 17, 1910;
C. M. Cat. No. 61.4742.

Fig. 3. Anodonta grandis footiana (Lea). The form of the beach-pools, from Lake
Erie, beach-pools of Presque Isle, Erie, Erie Co., Pennsylvania, collected June 2, 1908;
C. M. Cat. No. 61.3651.

Fig. 4. Anodonta grandis footiana (Lea). Female of the form found in Presque
Isle Bay, from Lake Erie, Presque Isle Bay (flats near west end), Erie, Erie Co., Penn-
sylvania, collected July 8, 1910; C. M. Cat. No. 61.4733.

Fig. 5. Anodonta cataracta Say. Creek-form, about half-grown, from White
Clay Creek, Avondale, Chester Co., Pennsylvania, collected April 26, 1909; C. M.
Cat. No. 61.4167.

Memoirs Carnegie Museum, Vol. VIII.

Plate X.

Anodonta.

3G4

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATEIXI.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Anodonta cataracta Say. Gravid female of pond-form, of good size, from
Pond near Paper Mill, Lockhaven, Clinton Co,, Pennsylvania, collected August 24,
1909; C. M. Cat. No. 61.4173.

Fig. 2. Anodonta implicata Say. Normal specimen, rather swollen, from Timber
Creek, Newbold, Gloucester Co., New Jersey, collected by C. H. Conner; C. M. Cat.
No. 61.4442.

Fig. 3. Anodonta implicata Say. Young specimen of the river-form, from Dela-
ware River, Yardley, Bucks Co., Pennsylvania, collected August 24, 1908; C. M. Cat.
No. 61.3693.

Fig. 4. Anodonta ohiensis Rafinesque. Gravid specimen, rather large, from Lake
Erie, beach-pool of Presque Isle, Erie, Erie Co., Pennsylvania, collected June 2, 1908;
C. M. Cat. No. 61.3291.

Fig. 5. Anodontoides ferussacianus (Lea). Specimen of medium size, from Little
Shenango River, Greenville, Mercer Co., Pennsylvania, collected June 5, 1908; C. M.
Cat. No. 61.3294.

Fig, 6, Anodontoides ferussacianus huchanensis (Lea). Typical specimen of
medium size from Lake Erie, beach-pool of Presque Isle, Erie, Erie Co., Pennsylvania,
collected June 2, 1908; C. M. Cat. No. 61.3295.

Fig. 7. Alasmidonta (Alasmidonta) undulata (Say). Gravid female of normal
shape, from Conodoguinet Creek, Carlisle, Cumberland Co., Pennsylvania, collected
August 13, 1910; C. M. Cat. No. 61.4660.

Memoirs Carnegie Museum, Vol VIII,

Plate XL

A nodonta-A nadontoides-A lasmidonta.

366

MEMOmS OF THE CAKNEGIE MUSEUM.

EXPLANATION OF PLATE XII.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Alasmidonta (Prolasmidonta) heterodon (Lea). Adult male from the
Schuylkill Canal, Manayunk, Philadelphia Co., Pennsylvania, collected April 24, 1909;
C. M. Cat. No. 61.4250.

Fig. 2. Alasmidonta {Prolasmidonta) heterodon (Lea). Adult gravid female from
the Schuylkill Canal, Manayunk, Philadelphia Co., Pennsylvania, collected April 24,
1909; C. M. Cat. No. 61.4250.

Fig. 3. Alasmidonta (Decurambis) marginata (Say). Adult gravid female from
the Shenango River, Shenango, Mercer Co., Pennsylvania, collected September 27,
1909; C. M. Cat. No. 61.4270.

Fig. 4. Alasmidonta {Decurambis) marginata susquehannce Ortmann. Male from
the set of types, from the Susquehanna River, Selinsgrove, Snyder Co., Pennsylvania,
collected August 14, 1910; C. M. Cat. No. 61.4079.

Fig. 5. Alasmidonta {Decurambis) varicosa (Lamarck). Adult female, from Prin-
cess Creek, Kunkletown, Monroe Co., Pennsylvania, collected June 14, 1910; C. M.
Cat. No. 61.4671.

Fig. 6. Strophitus undulatus (Say). Specimen supposed to be this species, from
Delaware River, Newbold, Gloucester Co., New Jersey, collected by C. H. Conner;
C. M. Cat. No. 61.4135.

Fig. 7. Strophitus edentulus (Say). Half-grown male, from a mill-race of Crooked

>•

Creek, Tioga, Tioga Co., Pennsylvania, collected August 19, 1909; C. M. Cat. No.
61.4154.

Fig. 8. Strophitus edentulus (Say). Adult gravid female, from Conneaut Outlet,
Conneautlake, Crawford Co., Pennsylvania, collected August 7, 1908; C. M. Cat. No.
61.3613.

Memoirs Carnegie Museum, Vol VIII.

Plate XII.

Alasmidonta-Strophitus.

368

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XIII.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. ElUpsaria fasciolaris (Rafinesque) . Gravid female from the Mahoning
River, Mahoningtown, Lawrence Co., Pennsylvania, collected August 4, 1908; C. M.
Cat. No. 61.3589.

Fig. 2. ElUpsaria fasciolaris (Rafinesque) . Inside of female shell, showing the
marsupial depressions, from the Shenango River, Pulaski, Lawrence Co., Pennsylvania,
collected July 19, 1909; C. M. Cat. No. 61.4128.

Fig. 3. ElUpsaria fasciolaris (Rafinesque). Form of Lake Erie, female, from Lake
Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected July 12, 1910; C. M. Cat.
No. 61.4748.

Fig. 4. Obliquaria reflexa Rafinesque. Male, largest found in Pennsylvania, from
the Ohio River, Industry, Beaver Co., Pennsylvania, collected October 5, 1909; C. M.
Cat. No. 61.4423.

Fig. 5. Cyprogenia stegaria (Rafinesque). Largest specimen found in Pennsylvania
in the Allegheny River, Godfrey, Armstrong Co., Pennsylvania, collected July 27, 1910;
C. M. Cat. No. 61.4760.

Fig. 6.- Ohovaria (Obovaria) retusa (Lamarck). The only male (dead shell) found
in Pennsylvania in the Ohio River, Industry, Beaver Co., Pennsylvania, collected
October 22, 1908; C. M. Cat. No. 61.3556.

Fig. 7. Obovaria (Obovaria) retusa (Lamarck). Gravid female, the only living
specimen found in Pennsylvania, in the Ohio River, Industry, Beaver Co., Pennsylvania,
collected August 29, 1908; C. M. Cat. No. 61.3555.

Fig. 8. Obovaria (Pseudoon) olivaria (Rafinesque). Male from the Ohio River,
Portland, Meigs Co., Ohio, collected September 22, 1910; C. M. Cat. No. 61.4776.

Fig. 9. Obovaria (Pseudobn) olivaria (Rafinesque). Female from the Ohio River,
Industry, Beaver Co., Pennsylvania, collected August 29, 1908; C. M. Cat. No. 61.3561.

Plate XIII

Memoirs Carnegie Museum, Vol. VIII

Ellipsaria-Obliquaria-Cyprogenia-Obovaria.

kf-

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. J

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I

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I

V

-Ji

.■ I

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fSA >■' '?*' ■•

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370

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XIV.

All figures are three-fourths natural size, a, lateral view; 6, dorsal view.

Fig. 1. Ohovaria (Obovaria) subrotunda (Rafinesque) . Probably male according
to shape, diameter 68 percent of length, from the Monongahela River, Charleroi, Wash-
ington Co., Pennsylvania, collected by G. A. Ehrmann; C. M. Cat. No. 61.2840.

Fig. 2. Obovaria (Obovaria) subrotunda (Rafinesque). Probably a female, judging
from the shape ; diameter 62 percent of length, from the Monongahela River, Charleroi,
Washington Co., Pennsylvania, collected by G. A. Ehrmann; C. M. Cat. No. 61.2840.

Fig. 3. Obovaria {Obovaria) subrotunda levigata (Rafinesque). Male, diameter 50
percent of length, from the Shenango River, Clarksville, Alercer Co.^ Pennsylvania
collected September 21, 1908; C. M. Cat. No. 61.3558.

Fig. 4. Obovaria {Obovaria) subrotunda levigata (Rafinesque). Female, diameter
52 percent of length, from the Shenango River, Clarksville, Alercer Co., Pennsylvania,
collected September 21, 1908; C. M. Cat. No. 61.3558.

Fig. 5. Actinonaias ligamentina (Lamarck). Male of medium size from the
Allegheny River, Kelly, Armstrong Co., Pennsylvania, collected October 4, 1907; C. M.
Cat. No. 61.3108.

Fig. 6. Actinonaias ligamentina (Lamarck). Gravid female of medium size from
the Allegheny River, Kelly, Armstrong Co., Pennsylvania, collected October 4, 1907;
C. M. Cat. No. 61.3108.

Fig. 7. Amygdalonaias truncata (Rafinesque). Male from the Ohio River, In-
dustry, Beaver Co., Pennsylvania, collected September 8, 1908; C. M. Cat. No. 61.3569.

Fig. 8. Amygdalonaias donaciformis (Lea). Male from the Missouri River, St.
Joseph, Buchanan Co., Missouri, collected by W. I. Utterback, November 1, 1913;
C. M. Cat. No. 61.6887.

Fig. 9. Amygdalonaias donaciformis (Lea). Female from the Missouri River,
St. Joseph, Buchanan Co., Missouri, collected by W. I. Utterback, November 1, 1913;
C. M. Cat. No. 61.6887.

Fig. 10. Plagiola lineolata (Rafinesque). Adult male from the Ohio River, Cook’s
Ferry, Beaver Co., Pennsylvania, collected September 12, 1908; C. M. Cat. No. 61.3565.

Memoirs Carnegie Museum, Vol, VIII.

Plate XIV.

■

Ohovaria-Actinonaias-Amygdalonaias-Plagiola.

F

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I

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372

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XV.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Plagiola lineolata (Rafinesque). Half-grown male from the Ohio River,
Cook’s Ferry, Beaver Co., Pennsylvania, collected September 10, 1908; C. M. Cat. No.
61.3565.

Fig. 2. Plagiola lineolata (Rafinesque). Adult gravid female from the Ohio River,
Cook’s Ferry, Beaver Co., Pennsylvania, collected September 12, 1908; C. M. Cat.
No. 61.3565.

Fig. 3. Plagiola lineolata (Rafinesque). Half-grown gravid female from the Ohio
River, Cook’s Ferry, Beaver Co., Pennsylvania, collected September 10, 1908; C. M.
Cat. No. 61.3565.

Fig. 4. Paraptera fragilis (Rafinesque). Male of Ohio-form from the Ohio River,
Industry, Beaver Co., Pennsylvania, collected September 23, 1908; C. M. Cat. No.
61.3549.

Fig. 5. Paraptera fragilis (Rafinesque) . Male of Lake Erie-form from Lake Erie,
Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected July 12, 1910; C. M. Cat.
No. 61.4807.

Fig. 6. Paraptera fragilis (Rafinesque). Female of Lake Erie-form from Lake
Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected July 12, 1910; C. M.
Cat. No. 61.4807.

Fig. 7. Proptera alata (Say). Male of Ohio-form, about half-grown, from Ohio
River, Industry, Beaver Co., Pennsylvania, collected August 30, 1909; C. M. Cat. No.
61.4108.

Memoirs Carnegie Museum, Vol. VIII,

Plate XV.

Plagiola-Paraptera-Proptera.

374

MEMOIES OF THE CAENEGIE MUSEUM.

EXPLANATION OF PLATE XVI.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Proptera alata (Say). Gravid female of Ohio-form, half-grown, from the
Ohio River, Industry, Beaver Co., Pennsylvania, collected August 30, 1909; C. M. Cat.
No. 61.4108.

Fig. 2. Proptera alata (Say). Gravid female of Lake Erie-form, nearly adult,
from Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected May 21,
1909; C. M. Cat. No. 61.4102.

Fig. 3. Toxolasma parvum (Barnes). Gravid female from Conneaut Outlet,
Conneautlake, Crawford Co., Pennsylvania, collected June 17, 1909; C. M. Cat. No.
61.4101.

Fig. 4. Eurynia (Micromya) fahalis (Lea). Male, according to shape, from Alle-
gheny River, Walnut Bend, Venango Co., Pennsylvania, collected September 6, 1908;
C. M. Cat. No. 61.3376.

Fig. 5. Eurynia {Micromya) fahalis (Lea). Female, judging from the shape,
from the Allegheny River, Walnut Bend, Venango Co., Pennsylvania, collected Sep-
tember 6, 1908; C. M. Cat. No. 61.3376.

Fig. 6. Eurynia {Micromya) iris (Lea). Male from Slipperyrock Creek, Wurtem-
berg, Lawrence Co., Pennsylvania, collected June 23, 1910; C. AI. Cat. No. 61.4827.

Fig. 7. Eunjnia {Micromya) iris (Lea). Gravid female from Little Beaver Creek,
Enon Valley, Lawrence Co., Pennsylvania, collected May 11, 1907; C. M. Cat. No.
61.2159.

Fig. 8. Eurynia {Micromya) iris novi-ehoraci (Lea). Alale from Conneaut Creek,
West Springfield, Erie Co., Pennsylvania, collected May 23, 1909; C. M. Cat. No.
61.4091.

Fig. 9. Eurynia {Micromya) iris novi-ehoraci (Lea). Female, judging from the
shape, from Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, collected July
7, 1910; C. M. Cat. No. 61.4831.

Fig. 10. Eurynia {Eurynia) nasuta (Say). Male from Lake Erie, Presque Isle
Bay, Erie, Erie Co., Pennsylvania, collected July 8, 1910; C. M. Cat. No. 61.4840.

Fig. 11. Eurynia {Eurynia) nasuta (Say). Female from Lake Erie, Presque Isle
Bay, Erie, Erie Co., Pennsylvania, collected July 12, 1910; C. M. Cat. No. 61.4840.

Fig. 12. Eurynia {Eurynia) recta (Lamarck). Male of medium size from French
Creek, Meadville, Crawford Co., Pennsylvania, collected September 1, 1908; C. M.
Cat. No. 61.3512.

Fig. 13. Eurynia {Eurynia) recta (Lamarck). Gravid female of medium size
from French Creek, Meadville, Crawford Co., Pennsylvania, collected September 1,
1908; C. M. Cat. No. 61.3512.

Memoirs Carnegie Museum, Vol, VIII,

Plate XVI.

Propter a- T oxolasma-Eurynia.

376

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XVII.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Lam.psilis luteola (Lamarck). Male from Mahoning River, Edinburg,
Lawrence Co., Pennsylvania, collected September 15, 1908; C. M. Cat. No. 61.3466.

Fig. 2. Lam.psilis luteola (Lamarck). Gravid female from Mahoning River,
Edinburg, Lawrence Co., Pennsylvania, collected September 15, 1908; C. M. Cat. No.
61.3466.

Fig. 3. Lampsilis luteola rosacea (DeKay). Male, deep water form, with very
regular growth-rests, from Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania,
taken at a depth of fourteen to sixteen feet, collected July 22, 1909; C. M. Cat. No.
61.4033.

Fig. 4. Lam.psilis luteola rosacea (DeKay). Male, form of the open shore, from
Lake Erie, outer shore of Presque Isle, Erie, Erie Co., Pennsylvania, from a depth of
ten feet, collected July 11, 1910; C. M. Cat. No. 61.4862.

Fig. 5. Lampsilis luteola rosacea (DeKay). Female, normal form of the bay,
from Lake Erie, Presque Isle Bay, Erie, Erie Co., Pennsylvania, taken from shallow
water, collected July 9, 1910; C. M. Cat. No. 61.4861.

Fig. 6. Lampsilis radiata (Gmelin). Alale from North Branch Susquehanna
River, Tunkhannock, Wyoming Co., Pennsylvania, collected August 21, 1909; C. M.
Cat. No. 61.4054.

Fig. 7. Lampsilis radiata (Gmelin). Gravid female from North Branch of the
Susquehanna River, Tunkhannock, Wyoming Co., Pennsylvania, collected August 21^
1909; C. M. Cat. No. 61.4054.

Fig. 8. Lampsilis ovata (Say). Young specimen, probably male, from French
Creek, Cambridge Springs, Crawford Co., Pennsylvania, collected September 13, 1909;
C. M. Cat. No. 61.4010.

Fig. 9. Lampsilis ovata (Say). Young specimen, probably female, but female
shape barely indicated, from French Creek, Cambridge Springs, Crawford Co., Penn-
sylvania, collected September 13, 1909; C. M. Cat. No. 61.4010.

Memoirs Carnegie Museum, Vol. VIII.

Plate XVII.

Lampsilis.

378

MEMOIRS OF THE CARNEGIE MUSEUM.

♦

EXPLANATION OF PLATE XVIII.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Lampsilis ovata (Say). Normal adult male from the Allegheny River,
Kelly, Armstrong Co., Pennsylvania, collected August 30, 1907; C. M. Cat. No. 61.2994.

Fig. 2. Lampsilis ovata (Say). Normal gravid female from the Allegheny River,
Kelly, Armstrong Co., Pennsylvania, collected August 30, 1907; C. M. Cat. No. 61.2994.

Fig. 3. Lampsilis ovata (Say). Male, strongly inclining in shape toward the var.
ventricosa, from the Allegheny River, Kelly, Armstrong Co., Pennsylvania, collected
July 5, 1909; C. M. Cat. No. 61.4007.

Fig. 4. Lampsilis ovata ventricosa (Barnes). Male, slightly inclining toward the
normal ovata, from French Creek, Cambridge Springs, Crawford Co., Pennsylvania,
collected September 13, 1909; C. M. Cat. No. 61.3996.

Memoirs Carnegie Museum, Vol. VIII,

Plate XVIII.

Lampsilis.

380

MEMOIRS OF THE CARNEGIE MSEUUM.

EXPLANATION OF PLATE XIX.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Lampsilis ovata ventricosa (Barnes). Quite normal gravid female from
French Creek, Cambridge Springs, Crawford Co., Pennsylvania, collected September
13, 1909; C. M. Cat. No. 61.3996.

Fig. 2. Lampsilis ovata ventricosa (Barnes). Typical male from the Shenango
River, Clarksville, Mercer Co., Pennsylvania, collected September 21, 1908; C. M. Cat.
No. 61.3404.

Fig. 3. Lampsilis ovata ventricosa (Barnes). Typical half-grown gravid female
from the Shenango River, Clarksville, Mercer Co., Pennsylvania, collected September
21, 1908; C. M. Cat. No. 61.3404.

Fig. 4. Lampsilis ovata canadensis (Lea). Male from Lake Erie, Presque Isle
Bay, Erie, Erie Co., Pennsylvania, collected May 21, 1909; C. M. Cat. No. 61.3982.

Fig. 5. Lampsilis ovata canadensis (Lea). Female from Lake Erie, Presque Isle
Bay, Erie, Erie Co., Pennsylvania, collected May 22, 1909; C. M. Cat. No. 61.3982.

Memoirs Carnegie Museum, Vol. VIII.

Plate XIX.

Lampsilis.

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. 13

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382

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XX.

All figures are three-fourths natural size, a, lateral view; h, dorsal view.

Fig. 1. Lampsilis fasciola Rafinesque. Alale from Shenango River, Shenango,
Mercer Co,, Pennsylvania, collected September 27, 1909; C. M. Cat. No, 61.4099,

Fig. 2. Lampsilis fasciola Rafinesque. Gravid female from the Allegheny River,
Warren, Warren Co., Pennsylvania, collected September 15, 1909; C. M, Cat. No.
61.4019.

Fig. 3. Lampsilis cariosa (Say). Very large male from the Chemung River, South
Waverly, Bradford Co., Pennsylvania, collected August 20, 1909; C. M. Cat. No. 61.4003.

Fig. 4. Lampsilis cariosa (Say). Alale of normal size from the Susquehanna
River, Duncannon, Perry Co., Pennsylvania, collected August 14, 1908; C. M. Cat. No.
61.3414.

Fig. 5. Lampsilis cariosa (Say). Gravid female from the Susquehanna River,
Duncannon, Perry Co., Pennsylvania, collected August 14, 1908; C. AI. Cat. No. 61.3414.

Fig. 6. Lampsilis ochracea (Say). Probably male, judging from the shape, hardly
half-grown, from an unknown locality in the Hartman collection; C. M. Cat. No. 61.4437.

Fig. 7. Lampsilis ochracea (Lea). Probably a female, judging from the shape,
hardly half-grown, from the Delaware River, Newbold, Gloucester Co., New Jersey,
collected by C. H. Conner; C. M. Cat. No. 61.4012.

Fig. 8. Lampsilis orbiculata (Hildreth). Half-grown male from the Allegheny
River, Godfrey, Armstrong Co., Pennsylvania, collected October 8, 1908; C. AI. Cat.
No. 61.3504.

Memoirs Carnegie Museum, Vol. VIII.

Plate XX.

Lani'psilis.

384

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXI.

All figures are three-fourths natural size, a, lateral view; b, dorsal view.

Fig. 1. Lampsilis orhiculata (Hildreth). Half-grown female from the Allegheny
River, Godfrey, Armstrong Co., Pennsylvania, collected October 8, 1908; C. M. Cat.
No. 61.3504.

Fig. 2. Lampsilis orhiculata (Hildreth). Very large and much swollen gravid
female from the Ohio River, Industry, Beaver Co., Pennsylvania, collected September 23,
1908; C. M. Cat. No. 61.3500.

Fig. 3. Truncilla triquetra Rafinesque. Male from the Allegheny River, Kelly,
Armstrong Co., Pennsylvania, collected September 27, 1907; C. M. Cat. No. 61.2985.

Fig. 4. Truncilla triquetra Rafinesque. Gravid female from the Allegheny River,
Kelly, Armstrong Co., Pennsylvania, collected September 27, 1907; C. M. Cat. No.
61.2985.

Fig. 5. Truncilla- rangiana (Lea). Adult male from French Creek, Cochranton,
Crawford Co., Pennsylvania collected September 2, 1908; C. M. Cat. No. 61.3363.

Fig. 6. Truncilla rangiana (Lea). Half-grown female from French Creek, Coch-
ranton, Crawford Co., Pennsylvania, collected September 2, 1908; C. M. Cat. No.
61.3363.

Fig. 7. Truncilla rangiana (Lea). Fully adult gravid female from French Creek,
Cochranton, Crawford Co., Pennsylvania, collected September 2, 1908; C. M. Cat.
No. 61.3363.

Memoirs Carnegie Museum, Vol. VIII.

Plate XXI.

Lampsilis-Truncilla.

••I*

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'1

I . Ilia Court Held at
. ' ; t) for the District

• 1 76. CEUMEmu 90

. lia Court Held for

i . _ at Augusta Town

(now Washington, Pa.), and afterward on the
Andrew Heath Farm near West Elizaheth,
1776-1780. Ceumeine 2.26

23. Idinute or Order Book of the Virginia Court

Held for Ohio County, Virginia, at Black’s
Cahin (Now West Liberty, W. Va.), &c.
Ceumeinb 1.60

24. The Becords of Deeds for the District of West

Augusta, Virginia, for the Court Held at Fort
Dunmore, &c. Gettmeine 1,76

25. Astropecten (?) montauus, &c. Douglass 10

26. Astrodon (Fleuroccelus) in the Atlantosaurus

Beds of Wyoming. Hatchee. (Out of Print.)

27. Osteology of tiie Idmicolae. Shupeldt 1.00

28. New Vertebrates from the Montana Tertiary.

Douglass 1.26

29. New Genus and Species of Tortoise from the

Jurassic of Colorado. O. P. Hat .10

30. Osteology of Oxydactylus. Peteesoht 1.00

31. Birds of Erie and Presque I^e. Todd .76

32. J. B. B[atcher. By W. J. Holland ...... .16

33. Tropidoleptus Fauna at Canandaigua Lake, etc.

Eaymond. {Out of print.)

34. Two Turtles from the Jn^th Biver Beds of

Montana. O. P. Hat. {Out of prmt.)

35. Preliminary List of Hemlptera of Western

Pennsylvania. Wietnee, {Scarce.) .60

36. Trilobltes of the Ohazy Limestone. Batmond. 1.00

37. Crawfishes of Western Pennsylvania. Oet-

mann LOO

38. The Geology of Southwestern Montmia. Doug-

lass .40

39. New Crocodile from the Jurassic of Wyoming.

Holland ..... .10

40. Procambarus, a New Snbgeuns of the Genus

Cambams. Oetmann .16

41. Fres^tation of Beprodnction of Diplodocus Car-

negei to the British Museum. Holland A5

42. Birds Collected near Mombasa by William Do-

herty. Holland 20

43. Hyoid Bone in Mastodon Am^canus. Hol-

land .10

44. Additions to Flora of Allegheny County, Pa.

Jennings .16

45. New Species of Kneiflia. Jennings .06

46. Occnxrmice of !lhl^ochJn palustris in Fa. Jen-

nings .06

47. New Species of Ibidinm. Jennings J.0

48. Agate Spring Fossil Quarry. Pbiebson 40

49. Two New Birds from British E. Afrio. ObeEt

HOLSEB .05

50. The Chazy Formation and Its Fauna. Bat-

mond L60

5L A New American Cybele. Nabeawat and Bay-

mono 45

52. Plastron of the Protosteginae. Wieland 46

63. New Species of Testudo from the IHocene, etc.

O.P.Hat .25

54. Miocene Beds of W. Nebraska and E. Wyoming

and Their Vertebrate Faunae. Pbiebson LOO

55. New ^peties of Loniceta ftom Pa. Jennengs. .05

56. Meryocochoems, etc. Douglass.

57. New Merycoidodonts. Douglass. (Nos. 56

and 57 sold together.) 1.00

58. Further Collections of Fishes from Paraguay.

Eigenmann, McAtee, and Waed 1.26

59. Cndetenmned Element in the Osteology of the

Mosasauridse. Holland 20

60. Gasteropoda of the Chazy. Batmond 1.35

61. Occurrence of Wynea Americana in Pa. Jen-

nings 06

62. Account of Pleistocene Fauna Discovered at

Frankstown, Pa. Holland 10

63. Vertebrate Fossils from the Vicinity of Pitts-

burgh, Fa. Case ..... 16

64. Elsenidm from the Black Biver Limestone near

Ottawa, Canada. Batmond and Naeeawat. . .20

65. Bhinoceroses from the Oligocene and Miocene

of North Dakota and Montana. Douglass.. .25

66. Fossil Horses from North Dakota. Douglass. .30

67. Oligocene Lizards. Douglass 20

68. The Type of Stenomylns gracilis. Peterson.. .25

69. New Species of Birds from Costa Bica, etc.

Caeeikee .10

70. Costa Bican Formicariidae. Cabbikeb .05

71. Vertebrate Fossils from the Port Hnion Beds.

Douglass .46

72. List of the L^idoptera of Western Pa., etc.

Engel L25

73. Fauna of Upper Devonian in Montana. Fossils

of Bed ShMes. Batmond 60

74. New Species of Procamelus from Upper Mio-

cene of Montana. Douglass 30

75. Sections of the Conemangh Series between

Pittsburgh and Latrobe, Pa. Batmond 35

76. List of the Unionldae of Western Pa. Oetmann 60

77. Cteologdcal BeconuaJssance in North Dakota,

Montana, and Idaho. Douglass 1.00

78. Botanical Survey of Presque Isle. Jennings. . 2.75

79. Sesqni-Centennial (Pittsburgh) Belies. Stewart 45

80. Dromomeryx, New Genus of American Bumi-

nants. Douglass 60

81. Fossils from Glacial Drift mid Devonian and

Mississippian near MeadviUe, Pa. Millard. . .10

82. New Species of Helodus. Eastman .05

83. Charles Chauncey Mellor. Holland .15

84. Beports of Expedition to Britidi Guiana, etc.

Eigenmann 60

85. Beporfai of Expedition to British Guiana, etc.

Dubbin .26

86. Odonata of the Neotropical Begiou, Ifodnsive

of Mexico and Central America. Calvert. . . 2.25

87. Deinosnehns hatcheri, a New Gmms and £^;iecies

of Crocodile. Holland 60

88. B^orts on Expedition to British Guianay etc.

Blosseb .16

89. Description of Some New Titaaotheres from

tiie Uinta. Douglass ^

90. Annotated List of the Birds of Costa Bica In-

dading Cocos Idand. Cabbisxb 3.00

9L Geology of the Coast of the State of Alagdas,

BrazU. Bbannee .40

92. Fossil Fishes from the Bitnminons Shales at Bi-

adio Doce, Alagdas, Brazil. Jikdan 65

93. Ordovldaa Trilobltes^ No. IL Asapbidm from

the Beekmantown. Batmond 65

94. Ordovidan Trilobitef^ No. lEL Batmond &

Nabxawat

65

95. Ordovician Trilobites, No. IV. Eaymond .... .40

96. Fishes from Irkutsk, Siberia. Jordan and

Thompson 30

97. South American Tetrigidse. Bruner 1.00

98. Fauna of the Allegheny and Conemaugh Series

in Western Pa. Raymond 30

99. Ichthyological Survey About the San Juan

Islands, Washington. Starks 75

100. New Species of Pygidium. Eigenmann .10

101. The Brachiopoda and Ostracoda of the Chazy.

Raymond .50

102. A New Camel from the Miocene of Western

Nebraska. Peterson .15 -

103. Skeleton of Stenomylus hitchcocki, etc. Peter-

son .15

104. Skeleton of Diceratherium cooki. Peterson . . ' .15

105. Carnegie Museum Expedition to Central South

America, 1907-1910. Holland 15

106. Report Upon the Expedition of the Carnegie

Museum to Central South America. Hase-
MAN & Eigenmann 25

107. New Species of Fishes, etc., from Central South

America. Haseman 50

108. Annotated Catalog of Cichlid Fishes from Cen-

tral South America. Haseman 1.25

lOO. New Species of Fishes from the Rio Iguassu.

Haseman 65

110. Ornithology of the Bahama Islands. Todd &

Worthington .’ . .75

112. South American Acridoidea. I. Bruner .... 1.00

113. Species of Hasemania, Hyphessobrycon, and

. Hemigrammus. Ellis 25

114. New Characins in the Carnegie Museum. Eigen-

mann 30

115. Jurassic Saurian Remains Ingested within Fish.

Eastman 20

lie. Autograph Letter, of U. S. Grant to Edwin M.

Stanton. Holland .16

117. Albert J. Barr. By W. J. Holland 10

118. Seventeen New Neotropical Birds. Todd 25

119. Dr. David Alter, the First Discoverer of Spec-

trum Analysis. Holland 10

120. Two Mummy Labels. Allen 10

121. The Families and Genera of Najades. Ort-

MANN 1.00

122. Group of Stenomylins in the Carnegie Museum.

Peterson 25

123. Tertiary Fish-remains from Spanish Guinea.

Eastman 25

124. Plated Nematognaths. Ellis 66

125. New Species of Cambarus from the Isle of

Pines. OrtmaNn 15

126. Sedum Camegiei, a New Species of the Family

^ Crassulaceae, etc. Hamet 10

127. Two New Species of Fishes from Peru. Eigen-

mann r. ..... . .15

128. South American Locusts (Acridoidea). II.

Bruner .75

129. Revision of the Genus Chsemepelia. Todd 85

130. New Genus and Species of Abyssinian Rodents.

Childs Erick ; 20

131. New Titanothere from the Uinta. Peterson. . .20

132. Small Titanothere from the Lower Uinta. Peter-

son 10

133. Osteology of La

Shufeldt

134. New Rhynchoceph

135. Scales of South . X XSIIOS*

Cockerell .;.... 25

136. Skeleton of Platigonus Leptorhinus. Peter-

son ..... , . .10

137. Lichens Collected in the Thunder Bay District,

Ontario. R. Heber Howe, Jr. . . . . . . ... . ; , . .10

138. Preliminary List of Fossil Plants in the Roof

of the Pittsburgh Coal. Grier .......... .10

139. Undescribed Remains of the Uinta Titanothere

Dolichorhinus. Peterson -.15

140. Triassic Fishes Belonging to the Catopteridae , ^ :

and Semionotidae, Eastman .....' .16 -

141. Osteology of Promerycochoerus. Peterson . . . .40 '

142. Correction of a Generic Name. Peterson ...v, .05

143. Skull of Bison Crassicomis. Holland .... .10

144. Serrasalminae and Mylinaa. Eigenmann 60

145. Heads and Tails: Notes on Sauropod Dinosaurs.

Holland .' .16

146. Dipterus Remains from Upper Devonian of

Colorado. Eastman 10

147. Notes on Tropical American Tettigonoidea (Lo-

custodea). Bruner 1,66

148. New Species of Tortoise from Jurassic of

Utah. Gilmore .16

149. Fauna of Upper Devonian in Montana.

Haynes .46

150. New Sphagebranchus from Bahamas. Eigen-

mann 07

151. Marine Fishes from Colombia and Ecuador.

Wilson .15

152. Apareiodon, a New Genus of Characid Fishes.

Eigenmann .18

153. New and Rare Fishes from S. American Rivers.

Eigenmann 25

154. Three New Species of Characid Fishes. Eigen-

mann & Henn 12

155. The Species of Salminus, Eigenmann 7

156. South American Pceciliid Fishes. Henn 40

157. A New Species of Apatosaurus, Holland 7

158. Birds of the Isle of Pines. Todd 70

159. Reptiles and Amphibians of the Isle of Pines.

Barbour 20

160. Land and Fresh Water Shells of the Isle of

Pines. Henderson 20

161. Pelecypoda of the Chazy. Raymond 46

162. S. American Crickets, Gryllotalpoidea andAche-

toidea. Bruner 1.30

163. Preliminary Catalog of N. American Sphseri-

idae. Stbrki .... 76

164. Directions for Collecting Sphaeriidae and

Aquatic Gastropods. Sterki 10

165. Lepidoptera of the Isle cf Pines. Holland 60

166. Odonata of the Isle of Pines, Kahl 15

167. A Trip to Islands in Lake Erie. Goodrich. ... .15

168. Land-Shells of Islands at Western End of Lake

Erie, etc. Clapp .20

169. Orthoptera of the Isle of Pines. Holland &

Kahl ..... 16

Publiecdions of the Carnegie Museum, Serial No. 107.

MEMOIES

OF THE

OAENEGIE MUSEUM.

YOU VIIL NO. 2.

W. J. HOLLAND, Editor.

( FEB 26 192!

FOSSIL PLANTS FROM THE BEDS OF VOLCANIC ASH NEAR
MISSOULA, WESTERN MONTANA

By 0. E. Jennings

PITTSBUKGH.

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Osteology of Baptanodon. Gilmore

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Classification of Chalcid Flies. Ashmead

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41. Presentation of Reproduction of Diplodocus Car-

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MEMOIRS

05 THE

CARNEGIE MUSEUM.

VOL. VIII.

No. 2.

FOSSIL PLANTS FROM THE BEDS OF VOLCANIC ASH NEAR MIS-

SOULA, WESTERN MONTANA.

By 0. E. Jennings.

(Pl.vi'es XXII-XXXIII)

Introductory.

This paper deals with some fossil plants obtained by Mr. Earl Douglass from
the beds of volcanic ash in the vicinity of Missoula, Montana, and with a smaller
collection made by him near Winston, Montana. The specimens are now in the
Carnegie Museum.

The fossils from near Missoula were collected by Mr. Douglass from two places.
One of these places is termed on the labels ‘H^ocality 165” and is stated to be about
one and one-half miles north of Missoula, the collections having been made Sep-
tember 26-30, 1903. These beds are noted as being ‘'Several hundred feet thick
and are composed of volcanic ash, sandstone, conglomerate, etc. There are several
seams of coal.” Another label from the same place says: “In beds of volcanic
ash. Coal above and below.” The other station from near Missoula, and from
which the better part of the material came, is evidently near “Locality 165” and is
called on the labels “Locality 196.” The collection from Winston was made on
September 23, 1902, on the north side of Beaver Creek, northeast of Winston,
Montana. This latter collection is a small one, containing but two or possibly
three species.

The Missoula collections, with which this paper is mainty concerned, were

385

386 MEMOIRS OF THE CARNEGIE MUSEUM.

taken by Douglass from beds which he believed to be of Oligocene age.^ He states
that they “appear in the main to represent the Titanotherium and Oreodon beds
of South Dakota.” This horizon, as understood by most American paleontologists,
includes approximately the lower half of the White River formation.^ The
fossil plants in these deposits consist of impressions of leaves and of leafy twigs,
there being also a few impressions of fruits and leafless twigs. The Alissoula
specimens are in a very light colored, gra3’'ish- white, soft and friable rock, consisting
of a fine-grained volcanic ash, which was evident!}’' more or less stratified and lami-
nated."-^ It is generally believed that the dust was wind-borne and that it was
mainly deposited in freshwater lakes or other shallow basins. Douglass states
that “It does not ai)pear that the water was as a rule very deep. There are un-
doubtedly not only lake but marsh and river dejiosits. . . . We find nearly every-
where evidences of shallow water, such as rijiiile marks, bird tracks, plant remains,
shallow water molhisca, etc.” Rowc^ also states his belief that the ash fell in
freshwater lakes.

The Missoula specimens mostly preserve in considerable detail even the finer
venation of the leaf surfaces, and in a number of instances the outline of the whole
leaf is plainly evident. The Winston specimens are, however, in a harder, light
gray rock, which has slickenside surfaces developed at various angles and directions
and presents every appearance of having once been a slumping mass of very fine-
grained mud. In this material the fossils are unsatisfactory, the hardness of the
rock and its irregularity of fracture resulting in fragmentary specimens.

I have undertaken the study of these various collections more as a student of
modern systematic botany and ecology than as a paleobotanist, and it is barely
possible that my conclusions may in some instances differ from what might have
been those of a paleobotanist, trained as a stratigrapher. However, no sharp line
of distinction can now be drawn between the work of a paleobotanist on the one
hand and that of the student of modern botany and ecology on the other, and it is
plainly evident that each of these fields of study may yet receive many valuable
and enlightening contributions from the other.

In the preparation of the illustrations accompanying this article I have had
the able assistance of my wife, Grace K. Jennings, and to her is due quite largely

* Douglass, Earl. New Vertebrates from the Montana Territory. Annals Carnegie Museum, II,
1903, 145-199.

^ Willis, Baile3n Index to the Stratigraphy^ of North America. U. S. Geol. Surv., Prof. Paper
LXXI, 1912, p. 770.

^ Rowe, J. P. Some Volcanic Ash Beds of Montana. Univ. Montana Bull., XVII, 1903.

^ O/j. cit., p. 146.

® Op. cit., p. 12.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 387

the excellence of man}^ of the photographic reproductions. The photographs were
made with an ordinary 5 X 7-inch bellows camera, the specimens being usually
placed in direct sunlight at a low angle of incidence in order to give sharper contrast
and to show better the features of relief on the impressions. The photographs
showing parts of the specimens enlarged were made by supplementing the ordinary
lens of the camera with an enlarging lens which fits over the front of the regular
lens like a cap. To give greater contrast in these enlargements ordinary daylight
was supplemented with a rather strong desk dissecting lamp fitted with “daylight”
glass and by proper manipulation of this light the inequalities on the surface of the
leaf impressions were shown as highlight and shadow.

In the publication just referred to Rowe gives a short list of the plants collected
from beds of volcanic ash near Missoula, the list being as follows :

1. Sequoia Langsdorfii.

2. Sequoia, probably new species.

3. Glyptostrobus europceus.

4. Alnus.

5. Carpinus, probably new species.

6. Cornus or Viburnum.

7. Populus balsamoides (?)

8. Fruit near Chmchonidium.

9. Taxodium occidentalis.

10. Taxodium.

Accompanying Rowe’s report are three plates illustrating some of the fossil
plants from these beds. Plants shown on these plates are evidently the same as
some of those described in the present paper, as follows: Plate VI shows at the
left upper margin what is probably a piece of a leaf of Alnus Hollandiana Jennings,
a specimen of Populus Zaddachi Heer being shown in the middle of the plate,
while both Plates VI and VII show leaves of what I have described as a new species,
Betula multinervis. ^ The Taxodium-Y\]^Q sprays in Plate VIII are evidently the same
as the sprays in our material which I believe to represent one of the various types of
Sequoia Haydenii (Lesquereux) Cockerell.

The Glyptostrobus mentioned in Rowe’s list is probably the same as the speci-
mens in our collections which I believe belong to Sequoia Haydenii. The Carpinus
is evidently the same as my Betula multinervis] the Populus balsamoides (?) I take
to be Populus Zaddachi Heer and the Taxodium, like that shown in his Plate VIII,
at least, is probably Sequoia Haydenii.

II. Relations of the Fossil Flora from Missoula to the Nearest Eocene
AND Miocene Floras of the West.

The specimens collected by Mr. Douglass from near Winston appear to repre-
sent but two, or possibly three, species, as follows:

i

388

MEMOIRS OP THE CARNEGIE MUSEUM.

Equisetum insculptum Jennings
Equisetum sp. Vegetative buds
Aralia lo7igipetiolata Jennings

These species appear not to be reiiresented in collections described from other
localities and this fact, in connection with the wide geological ranges of the genera
Equisetum and Aralia, makes the collection of little value as a means of correlating
this flora with other ancient floras.

The siiecimens from the Missoula district rcjirescnt at least twenty species,
ten of which I have described as new and one of which recpiires a new name. These
twenty-oife species are enumerated in the following list, the numbers in iiarentheses
indicating from which of Douglass’ localities the specimens came:

Sequoia Haydenii (Lesquereux) Cockerell. (165, 196)

Sequoia ohlongifolia Jennings. (196)

Thuyopsis gracilis Heer. (196)

Sabina lingua’folia (Lesquereux) Cockerell. (196)

Typlia Lesquereuxii Cockerell. (196)

Cyperacites sj). (196)

Populus smilacifolia Newbeny. (165)

Populus Zaddachi Ileer. (165, 196)

Juglans pentagona Jennings. (165)

Betida multinervis Jennings. (165, 196)

Alnus microdontoides Jennings. (165, 196)

Alnus Hollanddana Jennings. (196)

Quercus approximeda Jennings. (196)

Quercus jlexuosa N ewberry . (165)

Quercus lourisinmlans Jennings. (196)

Ficus (?) priim/o/fa Jennings. (196)

/ lex furcinervis J ennings . (196)

Celastrus parvifolius Jennings. (196)

■Acer oregonianum Knowlton. (165)

Vaccinium pcdccocorymbosum Jennings. (196)

The flora rejiresented in the Missoula collections appears closely related to
that reported for the Florissant basin of Colorado.® Of the fifteen genera repre-

^ Among the more important titles consulted with reference to the Florissant flora were the following:

Kirchner. Trans. St. Louis Acad. Sci., VUI, 1898, pp. 161-198, Pis. 11-15.

Cockerell. LTniv. Colorado Studies, III, No. 3, 1906, 157-176. Bull. Am. Mus. Nat. Hist.,
XXIV, 1908, pp. 71-110, Pis. 6-10. Amer. Nat., XLIV, 1910, pp. 31-47.

Knowlton. Proc. U. S. Nat. Mus., LI, 1916, pp . 241-297, Pis. 12-27.

JENNINGS: FOSSIL PLANTS FEOM VOLCANIC ASH BEDS, WESTERN MONTANA. 389

sented in the Missoula flora all but two are also represented in the Florissant.
Three species, Sequoia Haydenii, Typha Lesquereuxii, and Sabina lingucefolia,
apparently occur in both floras and there are other species which a further study of
more complete and more abundant material might prove to be identical. Further
comparisons show that to a large extent the genera having more than one species
each in the two floras were largely the same. In the Missoula flora Sequoia,
Populus, Alnus, and Quercus were each represented by more than one species.

One of the fossils in the Missoula collection apparently represents the species
described by Knowlton from the Mascall beds of the John Day Basin, Oregon, and
in a number of respects the Missoula flora seems to bo rather closely allied to the
Mascall flora. Eleven of the Missoula genera occur in the Mascall and, while,
with the exception of the maple, perhaps none of the corresponding forms in the
two floras are identical, some of them are rather closely similar. Of the genera
reported as having two or more species each, the Mascall flora has Sequoia (with
2 or 3), Quercus (5 or 6), Celastrus (2), and Acer (8), as against Sequoia (2), Populus
(2), Alnus (2), and Quercus (3) in the fossil flora at Missoula.

Another of the floras of the John Day Basin reported by Knowlton’^ and of
interest with reference to the Missoula collections is that from what are regarded as
upper Eocene^ beds at Bridge Creek, Oregon. Comparing the Missoula flora with
that of Bridge Creek it appears that seven of the fifteen genera of the former are
represented also in the latter. Of the genera of the Bridge Creek flora represented
by more than one species there are Sequoia (with 2 species), Juglans (3), Betula (4),
Alnus (4), Quercus (8), Ficus (1), Acer (1). The Alissoula flora has Sequoia (2),
Populus (2), Alnus (2), and Quercus (3), so that, considering the relative percentage
of species represented by the leading genera together with the number of genera
common to the two floras, it would appear that the flora from Missoula is about as
closely related to the Bridge Creek flora as to that of the Mascall beds.

Knowlton® has reported eighteen genera of plants among the fossils collected
in the Paj^ette formation (regarded by Knowlton as Upper Eocene)^® from the

^ Knowlton. Fossil Flora of the John Day Basin. U. S. Geol. Surv. Bull., CCIV, 1902, pp. 19,
89-93, 106-108, 113. Op. cit., pp. 17, 89-93, 103-105, 113.

* Merriam, J. C. Significant Features in the History of Life on the Pacific Coast, in Nature and
Science on the Pacific Coast, 1915, pp. 88-103. See also A Contribution to the Geology of the John Day
Basin. Bull. Dept. Geoh, Univ. Cal., TI, 1901, 269-314, 285-287, 290-299; also Merriam, J. C.,and
Sinclair, W. J. Tertiary Faunas of the .John Day Basin. Bull. Dept. Geoh, Univ. Cal., V, 1907, p. 173.

® Knowlton. The Fossil Plants of the Payette Formation. U. S. Geol. Surv. Ann. Rpt., XVIII,
Part III, 1898, pp. 721-736, Pis. 99-102.

Knowlton. Succession and Range of Mesozoic and Tertiary Floras, in Willis and Salisbury,
Outlines of Geologic History, Chapter X, 1910, pp. 200-211.

390

MEMOIRS OF THE CARNEGIE MUSEUM.

fossil lake beds of the Snake River in western Idaho. Eight of these genera are
common to the Payette and Missoula collections. Of the Payette genera five are
represented by more than one species each {Sequoia, Myrica, Populus, Betula, and
Quercus) as against four such genera in the Missoula collections {Sequoia, Populus,
Alnus, and Quercus). While not identical in the two floras a number of species
are here also represented by similar forms.

The Lamar flora described by Knowlton^^ from the Yellowstone National Park
has listed for its lower member (“Fossil forest a”) eight of the fifteen genera which
I have recognized in the Missoula collections, and there are at least five or six
species closely similar in these two floras.

From what he regards as the true Green River formation “excluding Florissant
and Elko Station/’ CockerelP^ has compiled a list of the plant genera represented.
A comparison of this list with the list of Missoula genera shows nine genera to be
common to the two floras.

The following table presents in a more compact form a list of the plant genera
represented in the Missoula collections together with the occurrences of the genera
in the other floras discussed in the preceding pages :

Table of the Genera Represented in the Collections of Fossil Plants from the Beds of Vol-
canic Ash near Missoula, Montana, showing also the Occurrence of these Genera in a

Number of other Fossil Floras.

Regarded as Upper Eocene.

Middle

Miocene.

White River.
Missoula, Mont.

Lamar.
Yellowstone
National Park.

Green River.
Wyoming and
Utah.

Payette.
Snake River,
West Idaho.

Bridge Creek.
Oregon.

Mascall.

Oregon.

Florissant.

Colorado.

Sequoia

X

X

X

X

X

X

Thuyopsis. . . .

Sabina

(x)

X

Typha

X

X

Cyperacites . . .

X

X

X

Populus

X

X

X

X

Juglans

X

X

X

X

X

X

Betula

X

X

X

X

Alnus

X

X

X

X

Quercus

X

X

X

X

X

X

Ficus

X

X

X

X

X

X

Ilex

X

X

Celastrus

X

X

X

X

Acer

X

X

X

X

X

X

V accinium . . .

X

“ Knowlton. Fossil Flora of the Yellowstone National Park. U. S. Geol. Surv., Monograph
XXXII, Part II, Chapter XIV, 1899, pp. 651-791.

Cockerell. The Fossil Flora of Florissant, Colorado. Bull. Amer. Miis. Nat. Hist., XXIV,
1908, p. 44.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 391

It is obviously unsafe in questions of comparison or correlation between
rather closely related floras to place much reliance on such evidence as may be
furnished by a small collection of fossil plants consisting mainly of leaf-impressions.
Conditions of deposition and fossilization are probably very rarely effective in
preserving a representative sample of a flora, unless the fossils collected are in
large numbers. As far as the indications go, it appears that the Missoula specimens
represent much the same kind of a flora as was preserved in the Florissant beds.
Yet differences between the Missoula flora and the Green River flora, as referred to
above, and generally regarded as Upper Eocene^^ in age,, is' by no means great.
The Florissant beds are now regarded as Miocene^^, Cockerell even advancing both
the Mascall and Florissant floras to middle Miocene.

As far as the genera are concerned collectively there has been comparatively
little change in the flora of the temperate regions of the United States since Floris-
sant times, and, allowing for a probably gradual cooling and a decrease in moisture
during the period between the Green River (Eocene) and the Florissant (Middle
Miocene), it appears that the changes in genera in this latter period were likewise
not great. Cockerell, comparing the Intermediate and Lamar floras on the one
hand and the plants “said to occur elsewhere or in the Eocene” on the other, shows
that these Yellowstone floras have “twenty-six plants specifically identical with
those of the basal Eocene” and, further, “The conclusion seems to be legitimate
that the Yellowstone Intermediate and Lamar florae are Upper Eocene, or at least
older than Miocene.” In his discussion of the records furnished by fossils as to
the distribution of the various floras through the different periods of time, Clements
says^® that “ the flora of the Oligocene was essentially that of the Eocene somewhat
reduced by deformation, and the plants of the Pliocene are practically those of the
Miocene, but with a striking reduction,” the reduction mentioned being due in
part to the reduction in the area in which sedimentation and fossilization were
taking place.

Of the floras referred to in the above discussion and in the table, the Bridge
Creek, Lamar, Payette, and Green River are now probably best regarded as late

Willis. Index to the Stratigraphy of North America. U. S. Geol. Surv., Prof. Paper LXXI,
1912, pp. 676, 758-760, 765.

Knowlton. A Review of the Fossil Plants in the United States National Museum from the
Florissant Lake Beds of Florissant, Colorado, With Descriptions of New Species and Lists of Type
Specimens. Proc. U. S. Nat. Mus., LI, 1916, pp. 241-297, Pis. 12-27.

Cockerell. The Miocene Trees of the Rocky Mountains. Am. Nat., XLIV, 1910, pp. 31-47;
Some American Fossil Insects. Proc. U. S. Nat. Mus., LI, 1916, pp. 81-106.

Clements, F. E. Plant Succession. Carnegie Inst. Wash., Publ. CCXLII, 1916, p. 352.

392

MEMOIRS OP THE CARNEGIE MUSEUM.

Eocene, the Mascall and Florissant as middle Miocene. Assuming this to be the
case it is interesting to group together the four upper Eocene floras for purposes
of comparison with the two Aliocene floras. Upon doing this it appears that of
the fifteen genera in our Missoula flora twelve occur also in the combined upper
Eocene floras and fourteen in the combined middle Miocene floras. This compari-
son does not show any great preponderance in favor of the Miocene, and it is
quite possible that climatic and other ecological conditions may, have brought
about similar groupings in the Missoula and Florissant floras somewhat out of
proportion to the actual systematic relationships of the floras existing at the time
these fossils were formed.

The Missoula flora jirobably occupied the shores and surrounding slopes of a
high mountain lake. Douglass says, however: ‘M'^here is doubt that the mountains
were as high during the White River epoch as at the present time.”^^ Cockerell
and other writers have referred to Lake Florissant as a mountain lake and some of
the differences between the Mascall and Florissant floras are thought by Cockerell
to be possibly due to the differences between a lowland flora, like that of the Mascall,
and one around a mountain lake, such as that of the Florissant.^® I am inclined
to believe that the similarity of habitat has brought about a similarity in the fossil
floras from Missoula and Florissant that m ay have obscured to a considerable degree
the actual difference of the two floras in point of time.

Two of the iilants represented in the Missoula collections {Thuyopsis gracilis
Heer, and Populus Zaddachi Heer) appear to be the species described and reported
by Heer from the Atane beds of Greenland. These beds were regarded by Heer^®
and some other authors”^ as Miocene, but they are more likely Cretaceous or early
Eocene.^" A considerable number of plants described from the arctic regions by
Heer and others from beds thought at that time to be Miocene appear farther south
during early and middle Tertiary times. This southward shifting of floral zones is
to be ascribed to corresponding climatic changes. Clements notes'^® that W
distinct cooling is indicated b}^ the flora of the early Eocene, and the usual accom-

Douglass. New Vertebrates from the Montana Territory. Annals Carnegie Museum, II,
190.3, p. 149.

Cockerell. The Miocene Trees of the Rocky Mountains. Am. Nat., XLIV, 1910, p. 37.

Heer, 0. Flora Fossilis Arctica I, 1868, pp. 98-99, and various other pages in this and other
volumes of the series.

Schuchert, C. Climates of Geologic Time. In Huntington’s The Climatic Factor. Carnegie
Inst. Wash., Publ. CXCII, 265. 1914.

Willis, Bailey. Index to the Stratigraphy of North America. U. S. Geol. Surv., Prof. Paper
LXXI, 1912, pp. 705 and 838.

Clements, F. E. Plant Succession. Carnegie Inst. Wash., Publ. CCXLII, 1916, p. 242.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 393

paniment of aridity is shown by the salt and gypsum beds of the Texas formations
of the period. The earliest Eocene flora, that described by Heer from Belgium,
indicates a temperate climate, characterized by Quercus, Castanea, Salix, Laurus,
Hedera, etc. Similar horizons are found in the lower Eocene of France and England.
At a later stage, palms, bananas, figs, cinnamons, etc., became dominant, indicating
a return to tropical conditions.’ With the period of “mountain making and vul-
canism” in the Oligocene there came another change to a cooler and drier climate.
Clements further notes“^ that the evidence indicates “that the Oligo-Miocene cycle
was marked by a general climate cooler and drier than that of the Eocene, and hence
by a differentiation of climates approaching that of today.” And, further, “So far as
dominants are concerned there appears to be little difference between the floras of
the Eocene and Miocene. The dominant tree genera appear to have been about
equally represented in both, and this is largely true of shrubs. . . . Thus, while
the flora remained largely the same, it must have undergone marked differentiation
and shifting as a result of the deformation and cooling which initiated the cycle.
The northerly climax zones must have been broadened as well as pushed to the
south. . . . Before the climatic effect of Oligocene deformation had disappeared
the deformation cycle of late Miocene and Pliocene had begun to culminate in the
Ice Age. Thus the shifting of the climatic zones took place only to the southward,
as well as downwards on the mountains.” The cooling during the Oligo-Miocene
cycle “from a tro]iical or subtropical climate to a warm temperate one over much of
the continent was permanent.”

Assuming, then, that during the period from the late Eocene up into the
Miocene there was, in general, a cooling and drying of the climate and a differentia-
tion of climatic regions and zones and, further, that during this time there was a
migration of northern plants southwards as well as down from the higher and
cooler habitats to the lowlands, it seems to me not unreasonable to believe that the
mountains of western Alontana during the Oligocene would have been populated by
the northern flora long before a similar region in central Colorado at a latitude of
about six hundred miles farther south. The Missoula region would likely have
been invaded by this northern flora long before the Alascall region lying to the
southwest in Oregon and probably on lowlands separated but little from the western
ocean either by distance or elevation. Unless excejition might be made in the case
of the Payette flora, I feel fairly certain that the Florissant was the only one of the
fossil floras discussed which approximated very closely the Missoula flora as to the

Schuchert. Op. cit.

Clements. Op. cit., pp. 364-366.

394

MEMOIRS OF THE CARNEGIE MUSEUM.

ecological conditions involved. I think that the similarity of the Missoula and
Florissant floras may actually be regarded as indicating a considerable time interval
during which similar stages in a southward migration of northern plants were
reached in the two floras. During the corresponding disappearance of the southern
plants the Missoula region would, of course, lose them before they would disappear
from the Florissant and, in this connection it is interesting to note that of the
southern element Florissant had Sapindus, Diospyros, Persea, Leuccena, Annona,
Ficus, and others, while only one such species, which I doubtfully determined as
Ficus, appears in the Missoula collection. All this leads me to believe that the
collections from Missoula represent an earlier period than do the Florissant collec-
tions, a period somewhere between the late Eocene and middle Miocene, and T
see no reasons for not accepting Douglass’ claim that the beds belong to the White
River formation and are of Oligocene age.

If we may accept the claim that there had come about a differentiation of
climates it would not be unreasonable to expect a considerable difference between
the flora of the Missoula White River and the Oligocene flora of the lower south-
eastern part of the United States, much as is the case today with the modern floras.
Berry has this to say with reference to the tropical character of the Oligocene flora
of the Catahoula sandstone of the Gulf Coastal Plain^^: “Finally, the facies of the
flora as a whole is that of the abundant floras found in the early Oligocene of
southern Europe, notably in Provence, France, in Tryol, and in Dalmatia and
Styria. Not only does it exhibit this parallelism with these European early Oli-
gocene floras, but when the genera are considered separately it appears that almost
without exception they have not been found in what are now temperate latitudes
in any beds younger than Oligocene.” In general this was a tropical flora and the
climate along the Gulf Coast at that time was evidently much more tropical than
now.^®“^‘ There was later a general cooling, possibly also a shifting of ocean currefits
bringing about a change from subtropical to cold-water conditions, with evidently
somewhat corresponding changes in the flora of the adjacent coasts. Berry has
shown by a study of the flora of the Calvert formation from Virginia and the Dis-
trict of Columbia that by the time of the middle Aliocene the climate in this region

Berry, E. W. The Elora of the Catahoula Sandstone. U. S. Geol. Surv., Prof. Paper XCVIII-M,
191G, pp. 227-243, Pis. 55-60.

Berrj", E. W. A Study of the Tertiary Floras of the Atlantic and Gulf Coastal Plain. Proc.
Anier. Phil. Soc., L, 1911, 311-315.

Dali, W. H. Contributions to the Tertiary Fauna of Florida, Part VI. Trans. Wagner Free
Inst., Ill, 1903, p. 1594.

JENNINGS: FOSSIL PLANTS PROM VOLCANIC ASH BEDS, WESTERN MONTANA. 395

was cooler, probably only warm-temperate.^* ^‘The Calvert flora was a coastal
flora of strikingly warm-temperate affinities, comparable with the existing coastal
floras of South Carolina and Georgia along the south Atlantic coast or with those
along the coast of the Gulf of Mexico from western Florida to eastern Texas.
The climate of the Chesapeake Miocene epoch, cooler undoubtedly than that of
the Apalachicola or preceding epochs, was neither cold nor cool temperate.”

It is evident that the Oligocene and middle Miocene floras of the middle and
southeastern Atlantic Coastal Plains were related ecologically, and, especially as to
the climatic aspect, were related to the Oligocene and middle Miocene floras of the
West in very much the same manner as are the modern floras of those regions.
I can see nothing in either the character or ecological relationships of the old south-
eastern floras that might serve as an argument against our belief in the Oligocene
age of the Missoula flora.

III. Ecological Conditions Indicated by the Fossil Flora from AIissoula.

As to the ecological relationships of the flora around the old Missoula basin,
it is interesting to compare the fossil flora with that, which in the case of a shower of
volcanic ash we might suppose would be rejiresented by plant materials imbedded
under conditions likeW to lead to fossilization in the basins of some of the lakes
now to be found in western Montana and Idaho. The plants preserved in the
Missoula fossil collections were practically all woody plants, evidently mostly trees,
and I have found particular!}" useful for the purposes of this comparison the biologi-
cal reconnoissances of Flathead Lake and several other smaller lakes in western
Montana by Elrod, and the more detailed studies of the forests of the Flathead
Valley by Whitford.*® To a considerable extent, also, I have relied upon first-
hand information gained during two weeks in the summer of 1915 which my wife
and I spent in botanizing around Lake Newman, along the Washington-Idaho
state boundary line about one hundred and sixtv miles west of Missoula.

Whitford’s classification of the vegetation in the Flathead Valley, western
Montana, in the midst of rugged mountains, shows it to consist of five main divi-
sions, as follows: I. Vleadow (hydrophytic) ; II. Englemann Spruce Forest (meso-

Berry, E. W., Physical Conditions indicated by the Flora of the the Calvert Formation. U. S.
Geol. Surv., Prof. Paper XCVIII-F, 1916, p. 66.

Elrod, M. J. A Biological Reconnoissance in the Vicinity of Flathead Lake. Univ. Mont. Bull.,
X, 1902, pp. 91-182, Pis. 18-46. (Biological Series III.)

Whitford, H. N. The Forests of the Flathead Valley, Montana. Bot. Gaz., XXXIX, 1905,
pp. 99-122, 194-218, 276-296.

396

MEMOIRS OF THE CARNEGIE MUSEUM.

hydrophytic) ; III. Western Larch — Douglas Spruce Forest (mesophytic) ; IV.
Douglas Spruce — Bull Pine Forest (meso-xerophytic) ; and V. Prairie (xerophytic).

Numbers I, II, III, and V of Whitford’s divisions contain more or less abun-
dantly species corresponding to those represented in the Oligocene flora of Missoula.
The Meadow (I) is usually along the end or border of a lake or else in a wet swampy
depression, and such a meadow with subsequent filling or draining will tend to be
invaded and eventually succeeded by Spruce Forest (II).

From the articles cited I gain that most of the meadows have, on the hummocks
or as a bordering thicket, the advance guard of the spruce forest in the form of
willows (Salix) and alders (Alnus). The thickets formed by these plants are
eventually entered by spruce {Picea Engelmanni) accomjianied by poplars {Populus
angustifolia and P. tremuloides) , birches {Betula papyrifera), and sometimes
Echinopanax horridum, Rhamnus alnijolia, and Cornus stolonifera. For the pur-
}ioses of the present discussion, the wet meadow and spruce forest will be treated
as one unit. They are both situated close to the water or are in the central part of
a wet de]3ression and, often the shore of the lake may be so dry that there is no
meadow, the spruce forest bordering the shore directly, with perhaps but a narrow
fringe of alders, willows, and a few other plants.

Three of the genera mentioned in the above paragraph were present also in
the Oligocene flora oi Missoula, these three genera including five of the fossil species.
Added to these five, should be the fossil species of Typha, Cyperacites, Vaccinium.,
and Ficus, which must have occupied either the wet meadow, its invading and
surrounding thickets, or the adjacent wet woods. Thus considered, the meadow or
its immediate borders or adjacent wet woods in the Flathead Valley would be quite
likely to contribute to the waters of the nearby lake leaves of representatives of at
least five of the genera found in the Oligocene fossil flora from Alissoula, while,
considered from the standpoint of systematic and ecological equivalents, similar
habitats around a lake in the Vlissoula district in Oligocene times might easily have
contributed at least nine of the twenty-one species found among the Missoula
' fossils.

Whitford’s third class, the one having the habitat next drier than the spruce
forest and thus usually the one next higher on the slope from the lake basin, is the
Western Larch-Douglas Spruce forest. Characterizing this forest are the larch
(Larix occidentalis) , Douglas Spruce {Pseudotsuga mucronatci) , and accompanying
them are the Lodge-pole Pine [Pinus Murrayana) , Lowland Fir {Abies grandis),
White Pine {Pinus nwnticola), Engelmann Spruce {Picea Engelmanni) , White Cedar
{Thuya plicata), and occasionally Abies lasiocarpa, Pinus ponderosa, Tsuga hetero-
phylla, and some other chiefly deciduous trees and shrubs.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 397

With gentle slopes rising from the lake basins this forest will usually be found
at some little distance from the open water, but on higher, drier, well-drained soil
on steeper slopes or benches along the lake shore, such forests occur close to the
open water. At Newman Lake near the Idaho-Washington boundary line patches
of this association occupy soil not over fifteen feet from open water and extending
down upon levels not over three feet above lake-level. Such a forest might easily
become a strong competitor with the meadow-spruce forest in contributing leaves
and other materials to a bed of volcanic dust accumulating in the lake close by.
The old ecological equivalent of this habitat probably furnished for fossilization
in the Oligocene lake-bed at Missoula the two species of Sequoia, the Thuyopsis
(represented in our Northwest now by Thuya), the maple {Acer), the walnut
{Juglans), and perhaps some of the oaks (Quercus).

On sandy or rocky habitats or on diy uplands and slopes, where not too cold,
the next drier habitat to that occupied by the Larch-Dougias Spruce forest is that
of the Douglas Spruce-Bull Pine forest. On rocky or dry sandy soils around New-
man Lake this forest comes down practically to the water’s edge. Whitford^°'‘ notes
that: “If the outcrop is near a large body of water like Flathead Lake, Juniperus
scopulorum is one of the first trees.” The forest is mainly composed of the Yellow
Pine or Bull Pine (Pinus ponderosa) and Douglas Spruce. Such a forest around an
Oligocene lake-basin at Alissoula could thus have contributed readily from a
rocky cliff or promontory twigs of the Juniper {Sabina) or with the somewhat
warmer conditions indicated by the presence of the oaks and possibly a fig, the
Juniper might have come from a sandy shore or from sand dunes such as are now so
characteristically covered with the Bed Cedar {Juniperus virginiana) along the
shores of Lake Erie and the lower end of Lake Michigan.

Assuming that the climate in the Oligocene lake basin at Missoula was warmer
and probably drier {see p. 394) than that now prevailing in the Flathead Valley, it
appears probable that the habitat occupied by the Oligocene equivalent of the
Douglas Spruce-Bull Pine forest occupied a larger area, or at least crowded more
closely the more moist areas around the basin. I believe that this habitat is the
one which probably furnished the oaks, or at least most of them, and possibly also
the Holly {Ilex) and Celastrus, although modern species of both of these genera
also occur in more moist habitats. The Oligocene oaks were apparently xerophytic,
as indicated by the nature of the leaf impressions, and I take them to have been
comparable to the modern live-oaks, chestnut-oaks, and shingle-oaks so character-
istic of moderately xerophytic habitats in the eastern, southeastern, and south-
western parts of the United States.

Op. cit., p. 216.

398

MEMOIRS OF THE CARNEGIE MUSEUM.

As indicated in the preceding discussion, the fossils in the volcanic beds at
Missoula might readily have been contributed to the waters of an Oligocene lake,
in a climate somewhat warmer than now prevails in that region, by a series of
vegetational associations ranging from wet meadow to moderately xerophytic
oak-forests on rocky or sandy shores, all of these vegetational associations in close
proximity, at least here and there, to the waters of the lake.

IV. Remarks on the Fossil Flora from Winston, Montana

The collections from the consolidated mud from Winston apparently belong
to the same general age as the Missoula collections. As noted earlier in this paper-
there are but two species represented, a small Equiseiimi and a plant which I have
referred to Aralia. Neither of these plants offer much basis for correlation with
other fossil floras. Equisetum occurs in all of the late Eocene and middle Miocene
floras listed in the preceding table, excepting only Bridge Creek, and it, or the forms
referred to Equisetites, range from the Paleozoic up to modern times, so that its
presence means little as to the time relations of the beds in which it occurs. Aralia
occurs in the Lamar and Green River floras (late Eocene) and in the Florissant
(middle Miocene) . Its range from the upper part of the lower Cretaceous up to
modern times makes it, too, of little use for purposes of correlation. If there is
other evidence, as for instance, from animal fossils, for believing that these plants
were of Oligocene age I can see no objection to such a view from any evidence to be
furnished by the plant fossils. Ecologically the Equisetum and Aralia might readily
have been a part of the vegetation believed to have grown around the Oligocene
lake at Missoula; the Equisetum in shallow water with sandy bottom, the Aralia
in the thickets invading the wet meadow or in the moist woods.

V. Descriptions of the Species in the Oligocene Elora of Missoula,
Montana, and of a few Species from Winston, Montana.

1. Equisetum insculptum Jennings, sp. nov. (Plate XXII, Fig. 2.)

The specimen consists of a mould of a piece of stem about 3 cm. long, this
mould being mainly filled with the corresponding cast. The mould and cast repro-
duce in great detail the minute features of the outside of the stem. The stem was
about 3 mm. in diameter but was compressed in fossilization to about 4 mm. and
it had eighteen ridges about 0.5 mm. apart, these being quite prominent and each
having two distinct rows of papilla-like tubercles. In the rather deep and rounded
furrows there are fine but distinct and closely set silex ridges running crosswise and

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 399

there are longitudinal lines indicating the position of two rows of stomata. The

character of the impression suggests a species with a rather firm and strong stem-
wall, such as is the case in the living Equisetum hyemale Linnaeus.

The one specimen upon which this record is based comes from “Locality 139,”

near Winston. So far as I have been able to discover among the
various publications relating to fossil Equisetacem, there have been
no fossil species described in which the markings have been so
beautifully preserved as in this specimen. The most nearly related
species are, perhaps, Equisetum wyomingense, described by Les-
quereux from the Green River group of Wyoming, and the Equise-
tum studied by Knowlton from the Payette of Idaho, both prob-
ably Eocene in age. Among living species, Equisetum variegatum var.
Jesupi A. A. Eaton, which is probably a hybrid between E. variegatum
and E. hyemale^^ is very closely similar, having the double row of ca-
rinal tubercles, small stem, and cross-bands of silex in the furrows.

Fig. 1. Equise-
tum insculptum
Jennings. Sketch
to show details of
markings on the
stem sliown in
Plate XXII, fig. 2.
Enlarged about
fifteen diameters.

The genus Equisetum, or Equisetites, as most of its older representatives have
been called, is recorded from at least as far back as the Paleozoic. Excepting
possibly for a considerable diminution in size, these plants have changed but little
up to the present time and the various species are of relatively little stratigraphic
value.

As the Winston species seems not to be the same as any of the species before
described, I am designating it as new, the specific name insculptum being used for
it on account of the unusually distinct and bold markings on the outer surface of
the stem.

2. Equisetum sp. (Plate XXII, Fig. 1.)

This is a poorly preserved specimen from the Winstoa locality representing a
piece of a stem, about 17 mm. long and 3 mm. wide, at the apex of which is indicated
a cluster of three branches, only two of which are preserved with any degree of
distinctness. The most perfect of the branches consists of a short stem about 1 mm.
in diameter and 11 mm. long. This probably is a bud from the basal part of a
plant, just as may be found on some of the Equisetums of today, and such as have
been described several times in paleobotanical literature.

The specimen may represent a plant of the species to which I have given the
name Equisetum insculptum, found at the same place, but aside from a probable
similarity in size and the mere fact that they were taken from the same locality there
is no basis for such an assumption!

Holden, Ruth. The Anatomy of a Hybrid Equisetum. Am. Journ. Bot., II, 1915, pp. 225-237.

400

MEMOIRS OF THE CARNEGIE MUSEUM.

3. Sequoia oblongifolia Jennings, sp. nov. (Plate XXIII, Figs. 1 and la.)

Among the various fossils representing Sequoia in the White River material from
Douglass’ “Locality 196”, near Missoula, Montana, there are three specimens show-
ing fragments of leafy twigs, which can not be satisfactorily referred to any of the
various kinds of leafy twigs of Sequoia Haydenii, but apparently resemble rather
closely the specimens from Greenland described and figured by Heer as Sequoia
hrevifolia, as follows:^-” S. foliis oblongis, basi angustatis, adnato decurrentibus,
confertis, patentibus, planis, distichis, apice obtusis, infimis squanneformibus,
adpressis.”

As shown l)y Heer’s figures and later by Lesquereux^^ and by Knowlton,^^
the leaves of the White River plant were more strictly oblong, with straight sides,
and the base more abruptly rounded, while the leaf-blades were about one-half longer
and wider. The White River siiecimcns show the leaves about 7-10 mm. long
by 2-2.8 mm. wide, oblong, the apex rounded and apiculate. The midrib is fairly
strong, there being also faint furrows on each side of it and there are also faint indi-
cations of numerous fine longitudinal striae. The leaves were evidently quite thick
and stiff and the impressions show irregular cross-furrows and ridges, such as are
evident in dried specimens of many living species of Sequoia, Taxus, Podocarpus,
etc., having thick leaves. The base of the leaf is strongly decurrent, while the
leaves apparently spread distichously and at a wide angle, the blade often curving
enough to place the apex of the leaf at almost a right angle with the branch.

In general outline the leaves resemble some of the fossils described as Taxites
and Taxodium, but the quite strongly decurrent leaf base would appear to relate
it to Sequoia. It is also similar, possibly very close to some of the specimens referred
b}^ some authors to Sequoia Langsdorfd, but I have preferred to refer the Alissoula
specimens to a distinct species on account of the more decidedly oblong leaves
with the apex blunter than seems to be the case in S. Langsdorfd.

4. Sequoia Haydenii (Lesquereux) Cockerell. (Plate XXIII, Figs. 3, 4, 6; Plate
XXIV, Figs. 1, 2, 3; and Plate XXII, Figs. 3, 3a, 4, and 5.)

Hypnum Haydenii Lesquereux, Bull. U. S. Geol. and Geogr. Surv. Terr., I, 1875,

p. 383; Rept. U. S. Geol. Surv. Terr., VII, 1878, p. 44, PI. 5, figs. 14, 146.

Heer. Flora Fossilis Arctica, I, 1868, pp. 92 and 93, PI. 2, fig.s. 23 and 23/;.

Lesquereux, Tertiary Flora, U. S. Geol. Surv. Terr., VII, 1878, p. 78, PI. 61, figs. 25-27.

Knowlton. Flora of the Montana Formation. U. S. Geol. Surv., Bull. CLXIII, 1900, p. 27,
PI. 4, figs. 1-4.

JENNINGS; FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 401

Sequoia affinis Lesquereux, U. S. GeoL and Geogr. Siirv. Terr., Bull. I, 1875,
p. 384 (1876); Tertiary Flora, Rept. U. S. Geol. Surv. Terr., VII, 1878, p. 75,
PL 7; figs. 3-5 and PI. 65, figs. 1-4.

Glyptostrobus Ungerif Heer. Lesquereux, Cretaceous and Tertiary Floras,
Rept. U. S. Geol. Surv. Terr., VIII, 1883, p. 139, PL XXII, figs. l-6a.

Sequoia Haydenii (Lesquereux) Cockerell, Science, XXVI, 1907, p. 447; Pop.
Sci. Monthly, LXXIII, 1908, p. 122, text-fig.; Bull. Amer. Mus. Nat. Hist.,
XXIV, 1908, p. 78; Amer. Nat. XLIV, 1910, p. 32, fig. 3, p. 36.

The most abundant conifer in the material studied is a Sequoia which seems
to present a remarkable amount of variation in both size and form of leaf. After
much study I have decided to pursue the course adopted by Cockerell and Knowlton
with respect to the similar series of Sequoias found in the Florissant beds and to
regard these all as belonging to the plant described by Lesquereux as Sequoia
affinis and Hypnum Haydenii.

One common type of twig among these specimens is small and densely leafy,
the largest of this type (Plate XXII, figs. 3 and 3a) being about 7 cm. long, irregu-
larly pinnately branched, the leaves being mainly lanceolate or elliptic-oval,
ranging up to a size of about 3 to 4 mm. long by 1 mm. wide, tapering into a decur-
rent base about half as wide, and tapering above into an acute or somewhat obtuse
apex. The midrib is not prominent, often not showing at all. The leaves spread
at an angle of about 45°, being mostly somewhat recurved and being quite uniform
in size throughout the length of the twigs. These specimens match quite closely
the descriptions and figures of Sequoia Couttsice Heer,^° but the leaves from the
Montana collections are scarcely or not at all falcate and they are more spreading
than the Bovey Tracy specimens, as shown by Heer’s plates. Twigs of this type
are represented in material from Douglass’ “Locality 196 ” at Alissoula, but not from
Winston. This is evidently also the same plant as that occurring in some of the
Tertiary rocks of the Western United States and referred in the past to Glyptostrobus
europaus.

In association with the leafy twigs, but not attached to them, is a short twig
(Plate XXIII, fig. 4) at the apex of which are three cones, at least one of which,
and probably all three, belong to the twig. The cones are oblong, about 6 mm.
thick by 8 mm. long, rounded at the ends, the scales being 3-4 mm. long, with their
shank 4-angled, or perhaps some of them 3-angled, widening to about one and
one-half mm. at the apex, where they are capped by a thin flattened shield about

Heer. Fossil Flora of Bovey Tracy, Trans. Royal Soc. London, CLII, 1862, pp. 1051-1055,
PI. 59, figs. 1-19, PI. 60, figs. 1-45, and PI. 61.

402

MEMOIRS OF THE CARNEGIE MUSEUM.

2-2.5 mm. wide. The scales are loosely placed on the axis of the cone, the peltate
tops being only rarely in contact with each other and they are little or not at all
miicronate. These cones agree fairly well with the figures given by Saporta^® for
Sequoia Couttsice, but, as was also the case in the comparison of them with the figures
of the Bovey Tracy specimens, the Missoula cones are somewhat smaller and are
distinctly oblong.

In the Missoula matciial there is a small leafy shoot (Plate XXII. fig. 5)
about 12 mm. long, but with a rather thick central axis and clothed with leaves,
which range from scale-liLe organs less than 2 mm. long at the base to ascending
lanceolate bracts at the apex about 4-5 mm. long. The upper bracts apparently
form a cup or campanulate structure resembling the arrangement of the perigonial
bracts at the apex of a rigorous Poly trichum shoot. Indeed, the fossil suggests
at once a vigorous moss shoot. It appears, however, to be of the same nature as
the fossil described rather doubtfully by Lesquereux (l.c.) under the name
of Hypnum Haydenii. Lesquereux’s specimen has occasioned considerable dis-
cussion, Mrs. E. G. Britton and Dr. Arthur Hollick having referred it®’^ to some
conifer resembling Juniperus communis L., to young growing branchlets of which
the fossil shows considerable resemblance; while Cockerell {ll.ee., above) refers the
specimen to Sequoia affinis Lesquereux; and Knowlton, still later,®^ publishes it
as Juniperus? Haydeyiii (Lesquereux) Knowlton.

The Alissoula specimen is found to resemble very closely the short branchlets
which constitute the pedicels of the cones of Sequoia sempervirens Endlicher.
As compared with such branchlets on specimens of the Redwood in the Carnegie
Aluseum {Jennings, Mill Creek Valley above Muir’s Woods, Cal., 1915) the fossil
is found to have about the same size, the bases are similarly blunt and clothed with
short, thick, scale-like leaves, the apex having longer, more slenderly tipped bracts
arranged in campanulate fashion. In view of these similarities I am following
Cockerell in referring the moss-like twig to Lesquereux’s Sequoia affinis, which,
however, is required to be known as Sequoia Haydenii on account, unfortunately,
of the prior publication of the supposed moss.

The third type of specimen, which I am including under the name Sequoia
Haydenii, consists of rather densely complanately leaved twigs up to 12 cm. long
(Plate XXIII, fig. 6) and with leaves spreading collectively to a width of 7-30 mm.
The leaves become abruptly much shorter for a distance of 5 cm. or more at the base

Saporta. Annales des Sciences Naturelles, 5 Series, IV, 1865, PI. 2, figs. 2a and 2d.

Britton, E. G. & Hollick, A. Bull. Torrey Botanical Club, XXXIV, 1907, pp. 139-140.

Knowlton, F. Proc. U. S. National Museum, LI, 1916, pp. 249-250.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 403

of the twigs. The median leaves are linear-lanceolate to narrowly linear-oblong;
varying on different twigs from 5-17 mm. in length and from 1-2 mm. in width;
somewhat falcate, spreading at an angle of 50-70°. The apex of the leaf is usually
rather acute, sometimes more acuminate, while the base tapers to an oblique, some-
what thickened, and rather widely decurrent insertion. This latter character forms
perhaps the most easily observable distinction between this and the somewhat
smaller-leaved Taxodium dubium (Sternberg) Heer, to which there is a fairly close
resemblance in general aspect. Heer^® figures another character which apparently
distinguished Sequoia from Taxodium. In Sequoia there are strong cell-walls run-
ning from the midvein to the margin of the leaf and their course is often marked by
faint cross-lines on the outer surface of the leaf. In some of the Missoula fossils these
markings show quite distinctly. There is also a close resemblance to plates and de-
scriptions of Sequoia Langsdorfii (Brongniart) Heer which is a rather characteristic
Miocene species of Europe and has been reported from a number of localities in the
western part of the United States. It would seem not at all improbable that some
of the Western specimens reported as Sequoia Langsdorfii may belong to the same
species as do ours from Missoula.

Specimens of the complanately leaved type occur in the material from both of
Douglass’ localities 196 and 165.

5. Thuyopsis gracilis Heer. (Plate XXIII, Fig. 5.)

Thuyopsis gracilis Heer, Flora Fossilis Arctica, VII, p. 59, PL, XX, fig. 16; Schenck,
Palaeophytologie, ZitteFs Handbuch der Palaeontologie, II, 1890, pp. 322-323,
fig. 223.

The specimens which I have referred to this species consist of three small leafy
twigs, the longest of which is less than a centimeter in length. This longest twig,
shown in figure 5, matches so closely the figure of Heer’s species as to leave little
doubt as to the identity of the two plants. In our plants the leaves are appressed,
thick, smooth, evidently shining, those on the upper face of the twig being 3-4 mm.
long, 2-2.5 mm. wide, the lower two-thirds of only the margins being covered by
the overlapping edges of the adjacent pair of lateral leaves, the back of the leaf
having a median ridge about one millimeter wide and bordered on each side by
a slight furrow, which extends uniformly the whole visible length of the leaf.
The apex of the leaf is rather blunt, thick, and incurved. The lateral leaves are
apparently inserted a little lower on the twig, being about the same length as the
others, but somewhat narrower, the apex thickened and incurved. This approxi-
Flora Fossilis Arctica, I, 1868, PI. 2.

404

MEMOIRS OF THE CARNEGIE MUSEUM.

mation of the leaves into groups, almost whorls, gives the twig a peculiar jointed
appearance.

Cockerell has identified a fossil from the FlorissanC’ as Heyderia coloradensis
Cockerell, comparing it to the modern Libocedrus decurrens Torrey {Heyderia
decurrens Koch). This plant is evidently very similar to the plant from Missoula,
which I have called Thuyopsis gracilis Heer, but the specimens from Missoula show
strongly flattened shoots with short and wide leaves, the lateral leaves spreading
rather widely, and, so far as leafy twigs of such conifers can be taken to indicate
relationships, I think the evidence shows that they are Thuyopsis rather than
Libocedrus.

Thuyopsis gracilis was described by Heer from Greenland, from beds which he
regarded as Miocene, but which are now regarded as being older, perhaps Eocene
or even Cretaceous. The modern representatives of the genus consist of one species
native to Japan {Thuyopsis dolobrata Siebold & Zuccarini) and the variety nana
Siebold & Zuccarini {T. Icetevirens Lindley) native to China. So far as I know,
Thuyopsis gracilis has not before been reported on the North American continent.
The Montana specimens came from only one locality, Douglass’ “Locality 196” at
Missoula.

6. Sabina linguaefolia (Lesquereux) Cockerell. (Plate XXIII, Fig. 2.)

Glyptoslrobus europceus Heer. Lesquereux, Tertiary Flora. Kept, U. S. Geol.

Surv. Terr., VII, 1878, p. 74, PI. 7, figs. 1 and 2.

Widdringtonia lingucefolia Lesquereux, Cretaceous and Tertiary Floras, Rept.

U. S. Geol. Surv. Terr., VIH, 1883, p. 139, PI. 21, figs. 14 and 14a.

Sabina lingucefolia (Lesquereux) Cockerell, Univ. Col. Studies, III, 1906,
p. 175; Knowlton, Proc. U. S. Nat. Mus., LI, 1916, p. 249, PL 14.

These specimens consist of several fragments of leafy twigs, mostly less than a
centimeter long, some of them irregularly and closely pinnately branched. The
leaves are 4-ranked, thick, with a rather prominent dorsal ridge, which is especially
sharp at the apex. The branches are evidently complanately flattened, but the
smaller twigs appear almost as thick as wide. The facial leaves on the larger
twigs are about 1-1.5 mm. long, the lower half overlapped on the margins by the
lateral leaves, the exposed portion of the leaf rhomboid, almost as wide as long, the
apex acute and closely appressed, thick, with a sharp dorsal ridge. On these
larger twigs the lateral leaves are about the same size or perhaps a little longer,
their lower margins meeting over and below the lower part of the facial leaf for a
* Cockerell. Bull. Amer. Mus. Nat. Hist., XXIV, 1908, p. 78.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 405

distance often half that of the length of the exposed part of the facial leaf. These
lateral leaves are thick, ovate, somewhat bluntly acute, the tips spreading at an angle
of about forty-five degrees. On the larger twigs the successive pairs of lateral leaves
do not reach the next higher ones, this giving the twigs a jointed appearance, but
on the smallest twigs both the facial and lateral leaves are imbricate throughout.
On these smaller twigs the leaves are mostly less than 1 mm. long, scale-like, thick,
rather sharply dorsally carinate, there being some indication of a gland at about
the middle, the apex acutish, or on the lateral leaves often rather obtuse.

The specimens only occur in material from Douglass’ “ Locality 196, ” Missoula,
and they are all in mixture with detached Sequoia leaves and unidentifiable frag-
ments of wood, bark, etc., a general mixture of small floating fragments, such as
can often be seen stranded along more or less protected shores and banks during
quiet weather, or more rarely water-logged and forming a layer on the bottom of a
quiet pool.

The similarity of the foliage on these twigs to that of Chamcecyparis Ehrens-
wardi Heer is such that confusion might readily occur in attempting to state the
geological range of the two genera. Sabina lingucefolia is now known to occur in
the Florissant beds of Colorado, middle Miocene, and I would have little hesitation
in referring to the same species, or at least a very closely related one, the fragment
described and figured by Knowlton from Van Horn’s ranch, John Day basin, Oregon,
middle Miocene. The fossils from the White River beds differ scarcely at all from
impressions such as would be formed by the modern Rocky Mountain Juniper,
Sabina scopulorum (Sargent) Rydberg, occurring on foothills and bluffs throughout
the Rocky Mountains.

7. Typha Lesquereuxi Cockerell. (Plate XXIX, Fig. 4).

Typha latissima Lesquereux, The Cretaceous and Tertiary Floras U. S. GeoL
Surv. Territories, VIII, 1883, p. 141, PL 23, figs. 4, 4a. Not T. latissima
Al. Braun.

Typha Lesquereuxi Cockerell, Fossil Plants from Florissant, Colorado, Bulletin
of the Torrey Botanical Club, XXXIII, 1906, p. 307 ; idem, The Fossil Flora of
Florissant, Colorado, Bulletin of the American Museum of Natural History,
XXIV, 1908, pp. 77-110, Pis. 6-10.

The specimen consists of an impression of a leaf fragment about 1.5 cm. long
by 1.1 cm. wide, marked with fine longitudinal parallel veins about 1 mm. apart.

Knowlton. Fossil Flora of the John Day Basin, Oregon. U. S. Geol. Surv., Bull. CCIV, 1902,
p. 26, PI. 1, fig. 3.

406

MEMOIRS OF THE CARNEGIE MUSEUM.

Between these fine veins are traces of much finer veinlets, while the surface is
marked somewhat more strongly by fine transverse lines running from one vein to
the next. The leaf appears to have been fairly thick and smooth. Impressions
on the same piece of rock are probably those of stems but are indistinct and in-
definite.

According to the reports Typha appears first in the eastern United States in
the middle Cretaceous (Raritan and Magothy), next in the Eocene in Canada
(Paskapoo) and in Europe, then in the Oligocene of Europe and the Miocene of
Florida and Europe. The genus now contains about nine species well distributed
in fresh-water swamps in temperate and tropical regions.

Our specimen appears to be the same species as that figured by Lesquereux (l.c.)
and by Cockerell from the Florissant {l.c.), and, judging from the wide distribution
of some of the modern species, it is not unlikely that this species had a wide dis-
tribution in the Oligocene and mid-Tertiary swamps of western temperate America.

8. Cyperacites. (Plate XXIX, Fig. 3.)

There has been referred to Cyperacites a piece of a leaf measuring about 4 cm.
long by 4 mm. wide. The midrib is narrow and prominent, and there are indica-
tions of a less prominent vein on each side of it, as well as faint indications of other
longitudinal markings. The impression is practically uniform in width throughout
its length, and the indications are that the leaf was a leathery one with a glossy
surface, such as might be the case with coniferous leaves, as, for example, Podo-
carpus or Taxites Olriki, or with some of the species of the CyperacecB of the present
period, when growing in more or less xerophytic habitats. However, in the absence
of stems, flowers, or fruits, the identification of such a fragmentary specimen must
remain very uncertain. The specimen might with almost equal propriety be
regarded as representing a species of Podocarpus.

The specimen is from “Locality 196,” Missoula.

9. Populus Zaddachi Heer, (Plate XXV, Figs. 1,2; Plate XXVI, Figs. 1, la, 2,

3, 5, 6, and Plate XXIV, Fig. 6.)

Populus Zaddachi Heer, Flora Fossilis Arctica, I, 1868, pp. 98-99, Pis. VI, figs.
1-4, and XV, fig. 16; Lesquereux, The Tertiary Flora, U. S. Geological Survey
of the Territories, VII, 1878, p, 176, PL XXII, fig. 13; idem,. The Cretaceous
and Tertiary Floras, U. S. Geological Survey of the Territories, VIII, 1883,
p. 158, PI. 31, fig. 8.

The leaves are large, the blades being 5.5-16 cm. long, by 3-12 cm. wide,
ovate to broadly ovate, or the smallest ones more narrowly ovate, the base broadly

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 407

rounded to a rather narrow and shallow cordate sinus, the insertion of the petiole
having two well-marked glands at each side, or even with a glandular or thickened
rim on the outer side, as in some modern species of Populus, the upper part of the
leaf being somewhat acuminate with a rather obtuse apex. The leaves are seven-
to five-ribbed, the lowest pair of ribs weak and short, swinging back into the
broadly rounded basal curves, these, as well as all ribs, looping in camptodrome
fashion; the next upper pair in the seven-ribbed leaves leaving the insertion at
about right angles and extending well out to the margin of the blade in a broad
upturned curve; the pair next to the midrib are stronger, ascending at an angle of
about 45-35 degrees, approaching the margin at about two-thirds the distance
from base to apex of the leaf, and during the upper half or two-thirds of their
course throwing off from their lower side widely diverging and rather strong branches,
which, swinging around and forking, loop together in camptodrome fashion within
2-5 mm. of the leaf-margin. The veinlets from these latter loops usually also
form a series of much smaller loops from which faint veinlets run out to the small
crenate teeth. The uppermost pair of ribs is only about half as wide as is the
midrib, having thus about the same thickness as the lateral veins from the midrib
itself. The midrib is strong, about 1 mm. wide, sending out four to seven pairs
of widely- spreading, upwardly curving veins, either approximately opposite or
alternate, the strongest of these veins usually arising at about the middle of the
leaf or slightly below. Other fainter veins are thrown off at wide angles, some of
the finest of them at practically right angles, especially from the lower half of the
midrib. As compared with most broad leaves, the ultimate meshwork is rather
coarse. The texture of the leaves appears to have been rather thick and leathery
and the surface smooth, the larger nerves being deeply impressed on the upper
surface of the leaf and rounded but fairly prominent below.

In general shape the larger of these leaves resemble very closely those described
by Newberry as Catalpa crassifolia'^^ or later as Aristolochia cordifolia^^ . The
character of the teeth and the glandular insertion of the petiole can leave little
doubt, however, as to the propriety of placing these specimens in the genus Populus.
Berry'^'^ in connection with the description of Grewiopsis tennesseensis from speci-
mens from the Wilcox beds of Texas and Tennessee, calls attention to their resem-
blance to “the numerous forms from Greenland, Europe, and western North

Newberry. Annual Report, New York Lyceum of Natural History, IX, 1868, p. 56.

« Newberry. Illustrations of Cretaceous and Tertiary Plants, U. S. Geological Survey, IX, 1878, PI.
25, fig. 7; U. S. Geological Survey, Monograph, XXXV, 1898, p. 90, PL 39; PI. 40, fig. 7; and PI. 60, fig. 4.

Berry. The Lower Eocene Floras of Southeastern North America, U. S. Geological Survey,
Professional Paper XCI, 1916, p. 286.

408

MEMOIRS OF THE CARNEGIE MUSEUM.

America that are commonly referred to the genus Populus, as Populus arctica
Heer, Populus Zaddachi Heer, Populus cuneata Newberry (a variable and common
form of the Fort Union Eocene), Populus genetrix Newberry, Populus paleomelas
Saporta, or Populus glandulifera Heer.’’ He appears to doubt that these various
species belong to Populus and notes that “it is singular that the Arctic and early
American forms are palmately and not pinnately veined, like the modern species,
and present in a varying degree other distinctive features.” In connection with
this opinion attention should be called to the distinctly palmate ribbing of our
common modern Populus deltoideSy certainly just as distinctly five-ribbed as are
the five-ribbed leaves of the fossils at hand. In addition to the agreement in
venation, the presence of glands on the crenulations and at the insertion of the
petiole are further arguments for referring at least the Missoula fossils to Populus.
Sinnott and Bailey^® conclude among other things that “The primitive angiosperm
leaf was palmate in type, probably lobed, and was provided with three main bundles
which arose separately at the node;” and “This conclusion is based on evidence
from paleobotany, that the palmate leaf was more frequent in the Cretaceous and
Tertiary than at present.”

The petiole, when preserved, is slender, but apparently rather stiff and woody,
and distinctly enlarged at its base. The length of the petiole was 5.7 cm. in a leaf
with a blade about 12 cm. long. Another specimen, in which the blade of the leaf
was about 13 cm. long, had a petiole 6.2 cm. long.

The genus Populus is comparatively a very old one, its oldest known species
probably being Populus priniceva Heer, from the Kome beds of western Greenland,
which are presumably of Potomac age‘‘® (upper part of the Lower Cretaceous).
From this time on the number of species reported increases up towards the Eocene,
from which period there are perhaps forty species already reported from the United
States alone, many of these species resembling very closely some of the living species.
The range of Populus^ beginning in the far north in the Cretaceous, extended during
the latter part of the Cretaceous and in the Tertiary quite generally over the tem-
perate regions of the northern hemisphere. In western North America there have
been described from the Fort Union beds alone about as many species (25) as there
are now of living species of poplars in the whole world.

The range of Populus Zaddachi Heer is wide, the species having been reported

Sinnott and Bailey. Investigations on the Phytogeny of the Angiosperms, 5, American Journal
of Botany, II, 1915, pp. 1-23.

White and Schuchert. Cretaceous Series of the West Coast of Greenland, Bulletin of the Geologi-
cal Society of America, IX, 1898, pp. 365-367.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 409

from eastern Germany and western Greenland^^ and, in North America proper,
from Alaska, California, Colorado, Wyoming, Montana, and Dakota, from beds
variously ranging from the lowest part of the Eocene up to the Miocene.

The smallest of the leaves among the specimens are very likely those at the
apex of rapidly growing shoots, and they are much more narrowly ovate to even
lance-ovate than are the larger leaves, exactly as may be observed in Populus
balsamifera and other living species. Among the Missoula specimens is an im-
pression of a leaf-blade about 4 cm. long and 2 cm. wide (Plate XXVI, fig. 6)
which matches so closely one of the figures of Newberry’s Populus cuneata^^ that
Newberry’s figure might easily serve to illustrate, it It is not unlikely that imma-
ture leaves of a number of species of poplars, especially if preserved as fossils,
would be impossible or at least extremely difficult to refer to the proper species.
It is not at all improbable that some of the numerous fossil species of poplars repre-
sent such immature leaves, especially where the species have been described from
scanty material.

10. Populus smilacifolia Newberry. (Plate XXVI, Fig. 4.)

Populus smilacifolia Newberry, Annals of the New York Lyceum of Natural
History, IX, 1868, p. 66; idem, Illustrations of Cretaceous and Tertiary
Plants, U. S. Geological Survey of the Territories, 1878, PI. 14, fig. 5; idem,
Later Extinct Floras of North America, U. S. Geological Survey, Monograph
35, 1898, pp. 53-54, PL 29, fig. 5.

The specimen is from “Locality 165,” Missoula, and, although somewhat in-
complete at the apex, it agrees very closely with Newberry’s Populus smilacifolia,
from the Fort Union group of North Dakota.

The single specimen shows a rounded-ovate leaf from the cordate base of which
arise the midrib and two pairs of lateral veins, from the bases of the veins of the
lower pair of these arising, however, a much fainter and very short pair, so that the
leaf might be regarded as having three pairs of lateral veins. The lowermost
strong pair of veins spreads widely from the base of the leaf, curving upwards to
the margin at about half-way to the apex and sending off from the proximal side a
few widely spreading veinlets which curve upwards towards the margin. The
uppermost pair of veins, those next to the midrib, arise at a narrow angle with the
midrib and, curving slightly forwards, reach the margin but slightly below the apex.
In their upper two-thirds these veins give off short lateral branches, which curve out
Heer, O. Flora Fossilis Arctica, I, 1868, p. 98.

Newberry. The Later Extinct Floras of North America, U. S. Geological Survey, Monograph
XXXV, 1898, PI. 29, fig. 7.

410

MEMOIRS OF THE CARNEGIE MUSEUM.

and up towards the leaf-margin. So far as can be determined, the leaf-margin
appears to be but very slightly serrulate.

Populus smilacifolia as described and figured by Newberry does not have the
lowermost (third) pair of short and fine veins, as in the Missoula specimens, and
it may be possible that the latter represent only a form of the common Populus
Zaddachi Heer, the apex being but poorly preserved. The resemblance to Populus
smilacifolia is so close, however, that the specimen has been provisionally referred
to that species.

11. Juglans pentagona Jennings, sp. nov. (Plate XXIX, Figs. 1, la, 2, 2a.)

This species is based on the impression of a nearly complete leaflet about 6 cm.
long by 4 cm. wide, broadly ovate, rather abruptly rounded to a bluntly acute
apex, the base being apparently broadly rounded. The midvein is fairly strong;
secondary veins, about thirteen pairs, var3dng from alternate to opposite, the
lowermost arising from the midvein at a rather acute angle and then curving quickly
outward to a course at almost right angles to the midvein, then curving upward
toward the margin, the median and upper veins spreading from the midrib at an angle
of 25-30° and curving gradually upward towards the margin. All the veins unite
in wide camptodrome loops with the lower tertiaries from the next upper vein,
these loops usually running to within a millimeter or two of the leaf-margin and
giving off in turn finer loops reaching practically to the margin. The tertiaries from
the median and upper secondaries, especially, leave the secondaries at right angles
at distances of 2-4 mm. apart, fbrking and meeting in an alternate manner about
midway between the secondary veins, so as to form approximately pentagonal areas.
Within these meshes the finer veinlets form rather distinct polygonal meshes mainly
about 0.3-0. 7 mm. wide. The margin of the leaf is entire, but slightly undulate.
The lowermost secondaries are considerably closer together than are the median
and upper ones.

In another fragment of rock from this same locality is a leaflet which I take to
represent the same species as that described above. It is 8 cm. long by 2.2 cm.
wide, lanceolate, the base unequal, being rounded on the one side and tapering in
almost a straight line on the other side, the apex being rather slenderly acuminate.
The leaf -margin is entire. The midvein is strong, the secondaries being in about
seventeen pairs, the lowermost leaving at an acute angle, then curving more widely,
then again curving forwards towards the margin where they become camptodrome
with tertiaries from the next upper secondary. The median and upper secondaries
spread quite widely and towards the margin send off proximally strong tertiaries, or

JENNINGS: FOSSIL PLANTS PROM VOLCANIC ASH BEDS, WESTERN MONTANA. 411

they sometimes fork. Between these median and upper secondaries there are a
few intermediate shorter secondaries, which sometimes curve around and loop with
the main adjacent secondary at perhaps one-half to two-thirds of the distance from
the midvein to the margin.

The wider impression, the first of the fossils just described, is taken as the type
of the species, the specific name having been suggested by the more or less distinctly
pentagonal areas formed by the tertiary veins. The specimens are from Douglass’
‘^Locality 165,” Missoula, and are associated in the same blocks with Populus Zad-
dachi Heer, Betula multinervis Jennings, Sequoia Haydenii (Lesquereux) Cockerell,
and Alnus Hollandiana Jennings.

The genus Juglans has about twelve living species ranging from southeastern
Europe to eastern Asia, and in North America from the temperate regions south to
Mexico. The genus also occurs in the Andes of South America. About forty
fossil species have been referred to the genus, beginning rather early in the creta-
ceous (Dakota, Magothy, Raritan beds), the number of species increasing to be-
tween twenty and thirty in the Eocene, a few in the Oligocene, about forty reported
for the Miocene, and about thirty in the Pliocene. The genus was widely distrib-
uted over the northern hemisphere, especially during the Eocene, ranging from the
southeastern part of the United States to the middle and northwestern United
States, British Columbia, and Alaska. The distribution during the Aliocene was
apparently more southern (Florida, California, Brandon beds of Vermont, John
Day beds of Oregon), no occurrences having been noted farther north in America
than Oregon and Vermont.

Juglans pentagona is perhaps most nearly related among fossil plants to Juglans
crassifolia Knowlton, common in the uppermost John Day beds^® and regarded as
probably Oligocene, and to J. rugosa Lesquereux, reported from the Laramie and
Alontana formations (Cretaceous) and, more recently by Knowlton®*^ from the
Eocene (Raton Formation) of Colorado and New Mexico.

12. Betula multinervis Jennings, sp. nov. (Plate XXIV, Fig. 4; Plate
XXVII, Figs. 1, la, 16, Ic, 2; Plate XXVIII, Fig. 2.)

The leaf-blades vary from 4-9 cm. long to 2.5-5 cm. wide, being lance-ovare
to ovate-oblong or widely ovate, the petiole slender, about 1 cm. long, the leaf-base

Knowlton. Fossil Flora of the .John Day Basin, Oregon, U. S. Geological Survey, Bulletin
CCIV, 1902, p. 36, PI. 4, fig. 3.

“ Knowlton. Fossil Floras of the Vermejo and Raton Formations of Colorado and New Mexico,
U. S. Geological Survey, Professional Paper Cl, 1917, p. 293, PI. 112, fig. 4.

412

MEMOIRS OF THE CARNEGIE MUSEUM.

rounded or faintly subcordate, occasionally somewhat inequilateral, the apex
slenderly acute to long acuminate, and the margin of the leaf sharply doubly serrate
almost to the base. The tooth at the apex of each secondary vein appears larger
than the intermediate teeth, because the leaf -margin slopes inward and back-
ward from this tooth to a rather deep acute sinus in front of the next main tooth,
the intermediate teeth being not much smaller than the main tooth, but situated
successive!}^ lower down on the slope from it to the main sinus. The midrib and
veins are deeply impressed on the dorsal surface of the leaf, the veins being usually
in 15-16 pairs (13-18), close, almost straight, parallel; in the wider leaves giving off
toward the margin from one to four strong branches, which swing down and run
out into the intermediate teeth. The tertiary veins are rather prominent, mainly
running straight across from one secondary vein to the other and forming nearly
right angles with them, sometimes forking or irregularly anastomosing about halfway
between them. On the upper surface of the leaf the deeply impressed character of
the venation resembles the somewhat rugose surface, which occurs in some of the
living species of Ulnius.

The fruits associated with the leaves and probably belonging to dhe same
species, although not attached to the same twig with any of the leaves, occur only
in ‘‘Locality 165,” at Missoula. They are evidently narrowly oblong-cylindrical,
about 5 mm. thick by 1.5 cm. long, the scales being about 4 mm. long, their three
obtuse lobes reaching to about an equal height, the outer lobes about 0.8 mm. wide,
ascending at an angle of about 40° from a rounded sinus, the median lobe being
narrower. The main outline of the scale is basally rounded and then narrows rapidly
to an acuminate tapering portion about 1 mm. long.

The puzzling similarity between the leaves of species of Carpinus, Ostrya, and
Betula cannot but cause some uncertainty as to the generic position of the material.
It is quite generally stated that the teeth of Ostrya are simple and somewhat
smaller than are the more or less serrate teeth of Carpinus. The specimens show
rather fine teeth, which, as far as could be determined from the specimens from Mis-
soula can not be said to be serrate. Further, in the modern species of these genera
Carpinus has little or no indication of branches running out from the lower side of
the veins to form the secondary serrations, but this is quite characteristically the
case in our modern Ostrya virginiana. The specimens from Missoula agree in this
respect with Ostrya, rather than with Carpmus. However, this method of branch-
ing is also quite characteristic of some of the species of Betula, such as Betula lenta
of eastern North America, and a careful comparison of the specimens before me
with leaves of Ostrya and Betula reveals no diagnostic character by which the

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 413

fossil plant might be referred to one of these genera and not to the other. The
disposition of the species under Betula practically rests only upon the fact that there
were associated with the leaves some specimens of cones, which can be none other
than those of Betula, and as there are no leaves of other species among the fossils
which can be referred to Betula, it is assumed that the cones belong to the species,
the leaves of which are so abundant in the rocks at that place.

Impressions of these leaves occur abundantly in the collections from both of
the localities at Missoula, the most complete specimens being from “ Locality 196.”
The fruits are represented only from Locality 165.”

The genus Betula is reported first in North America from the Dakota group
in the Cretaceous, thence extending through the Montana and the Laramie into
the Eocene, where it is represented by more than a dozen species. In the North
American Miocene there are reported about a half-dozen species.

13. Alnus Hollandiana Jennings, sp. nov. (Plate XXV, Fig. 3; Plate XXIV,
Fig. 8; Plate XXVIII, Fig. 1; Plate XXX, Figs. 1, la,

Type; fig. 3, Cones.)

The leaves are variable in size, the blades of the largest being about 12 cm.
long by 5.5 cm. wide, elliptic to oval-elliptic, the apex being rather widely acute,
the base acuminately narrowed into a slender petiole, which in one specimen reaches
a length of 3.5 cm. The leaf-margins have small, low, somewhat crenate teeth,
which are tipped with a glandular or at least a thickened point, the teeth disappear-
ing along the basal 1-2 cm. of the margin. The midrib is rather strong, sending off
from nine to twelve somewhat upwardly curved parallel veins, the uppermost of
these more or less directly running out into the teeth, the greater number of them,
however, curving a little and with branches looping in camptodrome fashion
quite close to the margin and sending off minute branches which themselves ter-
minate in the teeth. Usually the base of the secondary vein bends down slightly
where it joins the midrib, thus forming a more acute angle. The tooth into which
the vein runs, especially in the upper part of the blade, or which most closely ap-
proaches the vein in the lower part of the blade, is somewhat larger than the inter-
mediate teeth, it having a height of 1-1.5 mm. There are three to five of these
intermediate teeth between each pair of the main teeth. The veinlets (tertiaries),
running across from one vein to the next, do so at about right angles to the veins and
are rather far apart (about 2-4 mm.), the enclosed areas thus being approximately
rectangular, and about two to three times as long as wide. The ultimate mesh work
is relatively quite fine, most of the veinlets visibly having their ends free. The leaves

414

MEMOIRS OF THE CARNEGIE MUSEUM.

appear to have been rather thin but yet stiff, leathery, and especially on the lower
side quite smooth and, perhaps, shining. The venation is impressed on the upper
surface of the leaf, even the finest meshwork showing on some of the impressions.
On the lower side of the leaf the venation is decidedly prominent, even to the finest
meshwork.

In some of the specimens the veins in the upper part of the leaf run almost
directly to the tooth, although sending off camptodromous branches towards the
margin, the tip of the vein which runs out into the tooth often being reduced to a
slender nervule not much stronger than those which run out into the intermediate
teeth. In general shape and also in venation there is considerable resemblance to
the terminal leaflet of some of the species of Hicoria and Juglans, but the resem-
blance to Alnus, such as Alnus rhomhijolia Nuttall, of California, or even of some
of the more acute leaves of forms of Alnus rugosa (DuRoi) Sprengel is closer.
In size and in the acute outline of the base and apex the leaves are likewise very
similar to those of the Mexican Alnus glabrata Fernald {See Pringle’s No. 8022,
from Tizapan, Mexico, 1899.)

The Missoula species is closely related to Alnus carpinoides Lesquereux from
Bridge Creek, Oregon, (Upper Eocene), but differs in that the leaves are more oblong
and less ovate than in the Oregon species. Practically the same characters distin-
guish the Missoula species from Alnus serrulata fossilis Newberry, also occurring
in the deposits of Bridge Creek, Oregon. The lower part of the leaf in the Missoula
specimens begins to narrow from about the middle of the blade and finally tapers
quite accuminately to the petiole.

There is a fine series of leaf impressions of this species from Mr. Douglass’
“ Locality No. 196 ” and a few from “ Locality 165,” many of them fragmentary, but
representing various sizes and considerable variation, principally with regard to the
apex, this occasionally becoming blunter, especially among the smaller leaves.
Among these smaller leaves it sometimes becomes difficult to make a satisfactory dis-
tinction between this species and the most acute forms among the larger leaves of
Alnus inicrodontoides with which latter species Alnus Hollandiana is frequently asso-
ciated in the same pieces of rock. Nevertheless, the majority of the specimens of
the two species show clearly as much difference as would be found in a similar asso-
ciation of specimens of leaves from different species of living alders.

This fine species is named in honor of Dr. William J. Holland, Director of the
Carnegie Museum, at whose request the study of the fossil plants from the Oli-
gocene beds of volcanic dust at Missoula was undertaken and which has led to the
preparation of this paper.

JENNINGS: FOSSIL PLANTS FEOM VOLCANIC ASH BEDS, WESTERN MONTANA. 415

14. Alnus microdontoides Jennings, sp. nov. (Plate XXIV, Fig. 7; Plate

XXX, Figs. 2 and 2a, Type.)

The leaves are 3.5-8 cm. long, and 2-3 cm. wide, widest somewhat above the
middle, the blade being narrowly obovate to broadly oblong-elliptic or even some-
what ovate, narrowed acutely to acuminately at the base, petioles slender, up to
13 mm. long, the apex widely rounded, the tip itself usually somewhat retuse
varying to obtuse or rarely acutish. The midrib is strong and deeply impressed
dorsally, ventrally standing out sharply and prominently. The secondaries are
also impressed dorsally and prominent ventrally, springing from the midrib at an
angle of about 45°, there being eight to eleven pairs, either alternate or opposite,
evenly spaced, parallel and straight to near the margin, then curving slightly for-
ward and usually ending in a tooth, but before doing so, branching and becoming
ver}'' slender, the branches being few, springing from the lower side of the vein and
curving out to the margin and there ending in teeth or variously anastomosing.
The tertiaries are spaced about 1-3 mm. apart, rather strong, distinctly impressed
above and prominent ventrally, springing from the secondaries at a wide angle and
curving slightly outward and running across to the next secondary, or forking and
joining with the forks of the adjacent secondary, the final mesh work being fine and
indistinct. The margin of the leaf is minutely serrate, there being from two to
four slightly smaller teeth between each pair of those teeth which terminate the
secondary veins. The teeth are sharply glandular- or callus-tipped and are
rather prominently directed forward.

In general appearance the leaves resemble rather closely the form of Alnus
rugosa known as variety serrulata (Aiton) Winkel {Alnus serrulata Willdenow).
They seem to bear much the same relation to this modern American plant that the
similar and approximately contemporaneous Alnus microdonta Saporta, from the
Armissan of southeastern France, bears to the modern Mediterranean Alnus
glutinosa var. denliculata (Meyer) Ledebour {Alnus oblonga Willdenow). In the
leaves of Alnus microdontoides the serrulations extend along the margins of the
leaves to within a centimeter or less above the the base, considerably farther than
Saporta figures for Alnus microdonta}'^

As has been pointed out in Gray’s Manual of Botany®^ there is considerable
intergradation, in the northern part of the range of Alnus rugosa, between forms
of that species and the equally variable Alnus incana, of somewhat more northern

Saporta, G. Etudes sur la Vegetation du Sud-est de la France a I’Epoque Tertiaire, Annales des
Sciences Naturelles, Botanique, Ser. 5, IV, 1865, p. 110, PI. 6, fig. 3.

“ Gray, Asa. New Manual of Botany, Seventh Edition, 1908, p. 337.

416

MEMOIRS OF THE CARNEGIE MUSEUM.

range. Specimens referable to Alnus rugosa show a greater variability than is
shown in the numerous leaf-impressions from the Missoula beds which I have
referred to Alnus microdontoides. During the first part of my studies on these
specimens the attempt was made to differentiate two species, one with a fairly
evenly serrate oblong-elliptic leaf, with a blunt or retuse apex and an acuminately
narrowed base, the other with a more plainl}^ doubly serrate leaf, relatively wider,
more acute at the apex and less acuminately narrowed at the base. Further
studies of material later found in another box with a large number of intermediate
specimens disclosed the fact that no separation of these extremes could be made
other than in a purely arbitrary manner.

Beginning in the upper Cretaceous, the genus Alnus attained in the Miocene
a general distribution through the arctic and northern North American and Eurasian
regions. Altogether there have been described about forty fossil species, mostly
from the Aliocene, although in the western part of the United States a number of
species have been reported from the Eocene, particularly from the Fort Union beds.
There are now recognized about thirty living species, these ranging well over the
northern hemisphere, and in America extending south to Peru.

15. Quercus laurosimulans Jennings, nom. nov. (Plate XXVIII, Fig. 3;

Plate XXXI, Figs. 3 and 3a.)

Quercus laurifolia Newberry, Proceedings of the United States National Museum,
V, 1883, p. 505; idem, The Later Extinct Floras of North America, U. S. Geo-
logical Survey, Alonograph XXXV, 1898, p. 76, PI. 59, Figs, 4 and 60, and
PL 60, fig. 3. Not Quercus laurifolia Michaux, Histoire des Chenes de
FAmerique, 1801, No. 10, PI. 17.

Newberry’s original description is as follows:

“Leaves petioled, lanceolate, 6 inches in length by l}/2 inches in width, equally
narrowed to the point and petiole; margins entire or faintly toothed or undulate;
nervation regular; midrib strong, straight, lateral branches, about ten pairs,
arching gently upward, terminating in the margins”.

The leaf-blade in the best Missoula specimen is between 10 and 11 cm. long,
by 4 cm. wide, elliptic lanceolate, rather slenderly acuminate at both base and apex,
and with a slender channeled petiole at least 2.5 cm. long, probably even longer.
The midrib is strong, rather wide (about 1 mm.), flat, sunken in the dorsal surface of
the leaf and sending off at wide angles about twelve pairs of slender but prominent
veins, which curve upwards towards the margin where the attenuate ends run almost
parallel to it and apparently form loops with the short branches from the next upper

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 417

vein. Between the main veins the midrib gives oft also short but prominent inter-
mediate veins forming practically right angles with it, these short veins forking and
disappearing at a distance of 4-6 cm. from the midrib. The veinlets from the
secondary veins are irregularly placed and form wide angles with the secondary
veins and finally anastomose to form a rather coarse meshwork with irregular
angles. The lowermost veins, even those in the attenuate base, leave the midrib at
about the same wide angle as those in the middle of the leaf, running quickly out
to the apparently slightly re volute margin. The margin is entire but slightly
undulate in places. The venation is mostly rather deeply impressed on the dorsal
surface of the leaf and prominent on the under surface. The extreme apex of the
leaf is broken off in the fossils at hand.

The name Quercus laurifolia having been used at an earlier date by Michaux
for the well-known Laurel Oak of the southeastern United States, it becomes
necessary to rename Newberry’s Tertiary species. The names lamina, laurifolia,
laurophylla, lauriformis and lauroides having been used for various species of fossil
or living oaks with leaves resembling those of some of the Lauraceoe, I have chosen
the name laurosimulans in order to preserve as nearly as possible the meaning of
the name used by Newberry for the fossil species.

The specimen first studied was referred to Laurus but further study led to a
decision in favor of Quercus. My reasons for referring the specimen to Quercus
rather than to Laurus are as follows: The rather numerous and prominent but
short veins given off from the midrib between the larger main veins and at approxi-
mately right angles to the midrib ; the rather strong branching and camptodromous
looping of the outer ends j of the veins; the veins in the attenuate base of the leaf
leaving the midrib at practically the same angle as those in the middle and upper
parts of the leaf; the impressed, but rather flat, dorsal surface of the midrib; and the
slenderer petiole.

The general outline of the leaf is very close, indeed, to that of Laurus princeps
Lesquereux, as figured from Corral Hollow, California. The resemblance to other
species of the Lauraceoe, such as Nectandra, is striking, as is also the resemblance to
the leaves of some of the living species of the Myrsinacece, such as Ardisia, Icacorea,
and Myrsine, but in these genera the midrib does not usually give off the short
intermediate veins such as appear so prominently in the Missoula specimens, or, if
present in the Myrsinacece, they are not at all prominent and, further, the petioles
are almost invariably quite short.

Newberry’s Quercus laurifolia was described from specimens obtained from
“Burned shales over lignite beds. Fort Berthold, Dakota,” now regarded as Fort

418

MEMOIRS OF THE CARNEGIE MUSEUM.

Union in age®^ (Eocene), thus not more distant from the Oligocene than many other
closely related species in the Tertiary,

About two hundred fossil species of oaks have been described, beginning in the
Cretaceous, during which period the genus was widely distributed in western Europe,
the United States, and in Greenland. In the Tertiary the genus occurred in Austra-
lia and quite abundantly in the northern hemisphere, in what are now temperate
regions, as well as farther north in Greenland and Alaska. In North America oaks
were particularly abundant in the Dakota and up through the Eocene and Miocene,
and in Europe they appear to have reached their maximum in the Oligocene and
Miocene.

Oaks with entire margins appear from the evidence thus far furnished by fossils
to have been relatively more numerous in the Tertiary than at the present time.
In the Oligocene and Aliocene there have been reported as oaks with entire margins
at least the following: Quercus neriifolia A. Braun, Q. elcena Unger, Q. Lyelli Heer,
Q. chlorophylla Unger, Q. Daphnes Unger, Q. elceomorpha Saporta, Q. lauriformis
Saporta, Q. soda Saporta, Q. areolata Saporta, Q. elliptica Saporta, and Q. simulata
Knowlton.

16. Quercus flexuosa Newberry. (Plate XXXI, Figs. 1 and la.)

Quercus flexuosa Newberry, Boston Journal of Natural History, VII, 1863,
p. 521; idem, The Later Extinct Floras of North America, U. S. Geological
Survey, Monograph XXXV, 1898, p. 74, PL 19, figs. 4-6.

The single specimen consists of the impression of part of a leaf, the base and
apex being missing. The impression shows the leaf to have been somewhat curved
and about 9 cm. long by 3.5 cm. wide, narrowly elliptic oblong, the basal portion
evidently narrowing inequilaterally. The margin has a few unequally spaced,
sharp, small teeth, these being directed forwards. The midrib is strong, the
secondaries fairly so, alternate, spaced about 5-9 mm. apart, arising at a wide
angle and swinging forwards, being branched towards the margin of the leaf and
these branches anastomosing in camptodrome fashion. Between the secondaries
the midrib sends off fine veins which anastomose with fine veins or their branches
from the secondaries, the meshes thus formed being pol3^gonal, with the sharper
angles and the longest dimension directed at about right angles to the midrib.

The specimen matches Newberry’s figures very closely as to size, outline, and
venation, although the inequilateral outlines in both Newberry’s figures and in

> 63 Merrill, G. P. Catalogue of the Type and Figured Specimens of Fossils, Minerals, Rocks, and

Ores. Part II. Bulletin of the United States National Museum, LIII, Part II, 1907, p. 283.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 419

the Missoula specimen suggests that the impressions may perhaps represent leaflets
of a compound leaf rather than a leaf of Quercus.

Newberry’s specimens were from near Bellingham, Washington, in rocks of
the Puget Sound group, thus possibly Oligocene, so that the species may have had a
general range over the region now separated into two floral regions by the Cascade
Mountains. The Missoula specimen was from Douglass’ “ Locality 165 ” and was
accompanied in the same block by Betula multinervis and Sequoia Haydenii.

17. Quercus approximata Jennings, sp. nov. (Plate XXVII, Figs. 3

and 3a, Type.)

The specimen consists of an impression about 6 cm. long from an oblong leaf
of a maximum width of 2.8 cm., the basal portion of the leaf being missing. The
leaf was pinnately veined, the midrib and secondaries being quite prominent, the
secondaries arising from the midrib about 7-11 cm. apart, but forming with the
midrib angles of but 20-30 degrees. Some of the veins are a little more widely
diverging at the very base, but immediately above this the veins all pursue straight
and parallel courses to the margin, where they end in rather low but sharply crenate
or forwardly-directed and evidently often slenderly spine-tipped teeth. Along
the margin of the leaf, between each pair of the main teeth, are from one to four,
usually two or three, similar teeth of about the same size as the main teeth. These
intermediate teeth are at the ends of fine tertiaries which swing off from the lower
side of the next upper secondary vein. On the specimen at hand one of the secon-
dary veins is strongly but acutely forked at about two-thirds of the distance from
the midrib to the margin of the leaf. The leaf was evidently distinctly leathery in
texture, the impression showing a smoothish surface on which the tertiary veins
are almost indistinguishable, being obscured by the very fine but unusually distinct
ultimate meshwork.

The specimen is from “Locality 196,” Missoula, associated in the same piece of
rock with fragments of two other leaves of probably the same species, and also with
Betula multinervis.

The species is nearly related to Quercus furcinervis americana Knowlton,
reported from the Miocene of the Yellowstone National Park (Lamar) and also
from the Eocene of the John Day beds of Oregon (Cherry Creek), and is perhaps
about as closely related to the following species reported by Heer from Greenland :
Q. Olafseni Heer, Q. platania Heer, Q. Steenstrupiana Heer, and Q. furcinervis
Rossmsessler. The species from Missoula differs from all of these, however, in
the much narrower angle between the secondary nerves and the midrib, and also
in that its tertiary veins are so much more obscure. The species clearly belongs to

420

MEMOIRS OF THE CARNEGIE MUSEUM.

the Chestnut-oak group, the straight, parallel, rather closely pinnate secondary
nerves, as well as the character of the teeth and the firm reticulate surface of the
leaf, all suggesting Castanea.

The specific name “ approximata” has been used for this species because of the
short distance between the secondary veins, this feature being due to the relatively
narrow angles formed by the secondaries with the midrib.

18. Ficus (?) prunifolia Jennings, sp. nov. (Plate XXXII, Fig. 1 and la, Type.)

The specimen consists of the impression from a piece about 7 cm. long from the
middle of a leaf about 7 cm. wide, pinnately veined, the secondaries being spaced
about 3-13 cm. apart, given off from the midrib at an angle of 50-55° and curving
gradually forward, in the outer part giving off strong branches which loop in
camptodromous fashion with the curved and attenuate end of the next lower secon-
daiy. From the midrib and from the lower side of the secondaries are given off
numerous fine but distinct veins, which pursue a course almost at 'right angles to
the midrib, and fork and anastomose to form polygonal, often diamond-shaped,
areas, the sharper angles and longest dimension of which lie cross-wise to the leaf.
Smaller and weaker camptodromous loops are formed closer to the leaf-margin by
branches from the main loops. So far as preserved the leaf-margin appears to be
entire, or at the most but slightly undulate. The venation appears to have been
rather deeply impressed and in texture the leaf was probably rather thick.

The specimen came from “ Localit}^ 196,” Missoula, and is on the same piece of
rock with imiiressions of Betula multinervis and Alnus Hollandiana. The generic
position of the sjiecies has been the cause of much perplexity. The rather striking
crosswise trend of the finer veins and their meshes is suggestive of species of Ficus,
Diospyros, Terminalia, various genera of the Ebenacece, Lauracece, and other
plants, but most of all, perhaps, it is suggestive of some of the larger leaved-species
of Prunus, such as P. alabamensis Mohr. This resemblance to Prunus, however,
is not so close, when the outermost course of the veins and the character of the
leaf-margin are also taken into account, so that the species seems best referred to
the genus Ficus.

In Ficus there have been reported })robably between two and three hundred
fossil species, beginning well down in the Cretaceous, thirty or more being reported
from the Dakota of the western i)art of the United States. During the Cretaceous
the genus was widely distributed, species being reported from Greenland, Europe,
North America, Australia, and New Zealand. In the Eocene there are about as
many reported, fifty or more; sixty or more from the Oligocene, mainly in western

JENNINGS: FOSSIL PLANTS FKOM VOLCANIC ASH BEDS, WESTERN MONTANA. 421

Europe; while about ninety species are reported for the Miocene, about twenty of
these being from the United States, and about twenty are reported from the Plio-
cene. The range of the genus has been extended by reports of its discovery in
Africa from the Oligocene and in Asia from the Miocene.®^

19. Ilex furcinervis Jennings, sp. nov. (Plate XXIV, Fig. 5, and Plate
XXXII, Figs. 2, 2a, and 2b, Type.)

The extreme apex and base of the leaf is lacking in our specimen. The im-
pression shows the leaf to have been obovate, probably about 8 cm. long, about 4
cm. wide, at the widest point, somewhat above the middle, narrowing rather rapidly
towards the base, apparently rounded or at least narrowing ver}^ rapidly towards
the apex. Margin entire in the lower half of the leaf, serrulate above, the teeth
being widely triangular, about 0.5 mm. high, sharply callus-tipped, this tip directed
forward and in the fossil showing a reddish-brown color. Midrib prominent,
sending off about twelve pairs of rather prominent and mostly alternate veins, which
leave at an angle of about 55-60 degrees, rather irregularly spaced, slightly curv-
ing upward, forking in their upper two-thirds or three-fourths, these forks them-
selves forking and their branches meeting to form irregularly diamond-shaped
areas extending to within 1-3 mm. of the leaf-margin. From the outer ends of
these diamond-shaped areas one or usually two nervules diverge, again forking
and looping in camptodromous fashion or disappearing into the minute teeth.
Usually it is the upper one of the two nervules leaving the apex of an area which
runs out into the tooth. In the middle of the leaf the tertiaries and the minute
ultimate meshwork are faint, suggesting that they were embedded in the tissue of
a thick leaf and are apparently rather irregular. The leaves were evidently not
only thickish but coriaceous and smooth.

In general outline the leaf was not unlike the wider leaves to be found on
vigorous shoots of our living southern Ilex Cassine Linnseus, and it also resembles to
a certain extent Ilex longifolia Heer,®^ both in venation and in that the margin is
toothed above. The general aspect of the leaf is that of the more obovate leaves
of Quercus imhricaria Michaux, our modern Shingle-oak, and there are resemblances
also to Quercus furcinervis anierica^ia Knowlton, reported for the Eocene and Mio-
cene.

The genus Ilex contains about 275 living species, mostly in North and South
America and temperate and tropical Asia, a few occuring also in Africa, Australia,

Berry, E. W. The Lower Eocene Floras of the Southeastern North America, U. S. Geological
Survey, Professional Paper XCI, 1916, pp. 82-83.

Heer, 0. Flora Eossilis Arctica, I, 1868, p. 124, PI. 48, figs. 3-6.

422

MEMOIRS OF THE CARNEGIE MUSEUM.

and Europe. More than one hundred fossil species have been reported, thirteen or
more of these having been accredited to the Upper Cretaceous from North America,
Greenland, and western Europe, and about fourteen in the Eocene from the eastern
and western United States, Alaska, Greenland, and western Europe. There are
twenty or more species reported for the Oligocene and more than fifty for the
Miocene.®® The occurence of Ilex in the Oligocene in the region of Missoula is
thus rather to be expected than otherwise.

The specimen in the collections of the Carnegie Museum consists of the obverse
and reverse impressions of a single leaf and it was found at ^‘Locality 196”, Missoula.
The specific name furcinervis has been chosen to indicate, as in the case of Quercus
furcinervis, the characteristic strikingly forked branching of the secondary veins.

20. Celastrus parvifolius Jennings, sp. nov. (Plate XXXI, Figs. 2 and 2a, Type.)

The single specimen shows an oblong-ovate leaf 11 mm. wide and evidently
about 3 cm. long, the apex being broken away. The base of the leaf narrows evenly
and rather acutely, being apparently sessile, but this latter feature can not be
stated definitely. The leaf is pinnately veined, the midrib being fairly strong, the
secondary veins being irregularly spaced, about four of them on each side of the
midrib, from which they spring at an angle of about 45 degrees. The secondary
veins curve gradually forward to near the margin of the leaf where they merge
into a series of camptodromous loops with branches from the next upper secondary
vein. The finer veins are quite irregular in spacing and direction, thus giving rise
to an irregular, although somewhat coarse meshwork. The leaf-margin, with
the exception of that of the cuneate base of the leaf, is rather finely serrulate with
low teeth, directed somewhat forward, there being usually a fine vein extending
into each of these teeth from the marginal camptodromous loops. The shape of the
upper part of the specimen suggests a rather slenderly tapering apex.

The specimen is from “Locality 196,” Missoula, and it perhaps most nearly re-
sembles Celastrus fraxinifolius Lesquereux, from Florissant, Colorado (Miocene),
and C. Lindgreni Knowlton, from the Payette of Idaho (Upper Eocene), although it
is smaller than either of these. The generic position is none too certain, the leaf
being suggestive of genera of the Rosacece, Leguminosae, Rhamnacece, and Ericacece.
Considering altogether the characters of leaf-margin, outline, and venation, par-
ticularly the method of looping and the finer meshwork, it seems best to refer the
plant to Celastrus.

See Berry, E. W. The Lower Eocene Floras of Southeastern North America, U. S. Geological
Survey, Professional Paper XCI, 1916, pp. 104-105.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 423

The genus Celastrus now contains over thirty species of shrubs, ranging
through southern and eastern Asia to Australia and America, mostly confined to
warm climates. The oldest known species {Celastrus arctica Heer) is reported
from the Cretaceous (Raritan and Magothy of New Jersey and Maryland, and the
Patoot of Greenland). About thirty species are reported from the Eocene,
ranging over England, the eastern and western United States, Alaska, and Greenland.
About the same number are reported from the Oligocene, evidently all European,
and fifty or more are reported from the Miocene of the United States (Virginia,
Colorado, Idaho, and Oregon), Europe, Asia, and Australia. Perhaps a dozen
Pliocene species are known.”

It is interesting to note in this connection that no species of Celastrus has been
reported heretofore from the Oligocene of America and that the geographically
nearest locality for the occurence of Celastrus in other horizons is that of the nearly
related Celastrus Lindgreni Knowlton in the Payette formation of western Idaho,
probably of late Eocene age.

21. Acer oregonianum Knowlton. (Plate XXXII, Fig. 3.)

Acer, fruits of, Lesquereux, Proceedings of the U. S. National Museum, XI,

1888, p. 15, PI. 6, figs. 2 and 3.

Acer oregonianum Knowlton, Fossil Flora of the John Day Basin, Oregon, U. S.

Geological Survey, Bulletin CCIV, 1902, p. 75, PI. 13, figs. 5-8.

Knowlton, l.c., says of this species: '' Fruits long and broad-winged, the axis
being evidently very thick and provided with numerous strong veins; nucleus
large, round, showing broad truncation where attached to the sister-fruit. . . . They
range in length from 3. 5-4. 5 cm. The wing is unusually broad, being not infre-
quently 1.75 cm. wide. It is filled with numerous strong veins which are given
off from the axis of the fruit in groups or bundles. . . . Abundant in Mascall beds
at Van Horn’s ranch and vicinity.”

The Missoula specimen consists of a part of a fruit, comprising the upper part
of the seed and a little more than 2 cm. of the wing. The bundles of strong veins
given off from the seed are quite prominent, these opening out into the membrane
of the wing at about right angles to the axis of the wing and forking from one to
three times. The widest part of the wing is about 14 mm.

This specimen is from “ Locality 165,” Missoula. As Knowlton has pointed
out, the fruit is very close to that of the modern Acer macrophyllum Pursh, found

See Berry, E. W. The Lower Eocene Floras of Southeastern North America, U. S. Geological
Survey, Professional Paper XCI, 1916, p. 105.

424

MEMOIRS OF THE CARNEGIE MUSEUM.

in the coastal region from Alaska to California, and now the only large native
maple-tree west of the Rocky Mountain region.

Acer now comprises about one hundred and twenty species, in the northern
hemisphere, mainly in temperate regions. Reports of the occurrence of the genus in
the Cretaceous are perhaps rather doubtful. More definite are the reports of its
occurrence in the Eocene (England) , while in the Oligocene various species occurred
in widely separated regions : Spitzbergen and southward in Europe to the Mediter-
ranean, Greenland, Alaska, and in the western United States. During the
Oligocene and Aliocene the species were especially numerous in western and central
Europe.

22. Aralia longipetiolata Jennings, sp. nov. (Plate XXXIII, Figs. I, la, 2,

2a, Type.)

Thematerial from “ Locality 139,’^ near Winston, Montana, contains numerous
fragmentary leaf-impressions, of which a number show the leaf-base and petiole,
but none of them show the complete apex. Although quite variable, the impres-
sions apparently all represent a species of Aralia.

The leaves were lance-ovate to oblong-ovate, reaching a length of 7 cm. or
more, the width being about half that amount. The petioles were slender, the
longest ones preserved being about as long as the leaves were wide. The leaf was
rather distantly pinnately veined, the secondaries being mostly six to eight on
each side of the midrib, the lowest opposite or essentially so, the median and upper
usually alternate. The veins form an angle of about 45 degrees with the midrib,
then curve gently forward, one occasionally forking in the outer third. The
secondaries are usually simple, but well out towards the margin they throw off at
wide angles from one to three tertiary veins, which curve around and, like the
attenuate tip of the secondar}^ end in low teeth, which are either sharply directed
forward or else are broadly rounded-crenate. The leaf-base was equally or some-
times unequally broadly rounded and shallowly cordate, most of the leaves having-
one or two jiairs of smaller more widely spreading secondary veins crowded into
the space below the lowest main jiair of secondaries. The appearance is almost that
of a sub-peltate leaf, remotely suggesting leaves such as those described at various
times under the names of Credneria, Aspidiophylhmi, and Protophyllum. The
tertiary veins are weak, rather closely spaced, and diverge from the secondaries
and from the midrib at almost right angles. They mostly fork and join midway
between the secondaries to form roughly pentagonal areas which are one to two
times as high as wide. As far as can be determined from the specimens, the apex

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 425

was acute or perhaps even slenderly acuminate. The impressions indicate that
the texture of the leaf was thin, like that of a shade-leaf.

Most of the fossil species of Aralia described have been of the type having
palmate veins and lobes, or else being palmately compound, but the Winston species
so closely resembles the modern Aralia racemosa L., that the writer has little hesi-
tation in referring it to the genus Ai^alia.

Broadly rounded crenate teeth, such as occur on the margin of a few of the
Winston fossils, are sparingly found in the modern Aralia racemosa, which rather
regularly has the petiole long and slender, the base unequal, the basal secondary
veins peculiarly crowded, and the leaf-blades thin, while the pattern of tlie vena-
tion, too, is much like that of the Winston fossils.

Although I can find no records of fossil Aralias ver}^ closely related to the species
from the Oligocene of Missoula, yet the present distribution of Aralia racemosa,
native to eastern North America, and closely allied species, such as A. cordata
Thunberg of Japan, and A. cachemirica Decaisne of the Himalayas, argues for a
Tertiary distribution of one or more ancestral forms of that type.

The genus Aralia now includes about thirty species of herbs and woody plants
ranging through temperate North America and Asia, and south from the latter
through the Malay Archipelago and into Australia. Alany of the fossil species
so reported are perhaps not Aralia, but belong to other genera of that family or
even to other families. However, taking the reports as they stand at the present
time, there are numerous species of Aralia described from the Cretaceous, perhaps
sixty altogether, beginning in the upper part of the Lower Cretaceous in Portugal
(Albian) and the eastern United States, and becoming numerous and practically
cosmopolitan in the Upper Cretaceous. From the Eocene about twent3^-five species
have been described, these being most numerous in the Fort Union of the western
United States and in the Paleocene of Belgium, while about an equal number have
been described from the Oligocene, all European, and more than half of these are
from the Sannoisian of southeastern France. About twentj'^-five species are recorded
from the Miocene, distributed through North America, Eurasia, and Australia.

Various species of Aralia having been recorded for the western part of the
United States from the Eocene and Miocene, it is to be expected that Oligocene strata
would also yield them from that region and, even though no closely related forms
may have been recorded from the Eocene or Miocene of the West, yet the present

See Berry, E. W. Aralia in American Paleobotany. Botanical Gazette, XXXVI, 1903, pp. 421-
428; idem, The Lower Eocene Floras of Southeastern North America, U. S. Geological Survey, Professional
Paper XCI, 1916, pp. 122-123.

426

MEMOIRS OF THE CARNEGIE MUSEUM.

distribution of Aralia racemosa and its allies would indicate that ancestral forms of
that species existed in the region, so that their occurrence in the Oligocene of the
Missoula district would be quite natural.

As the type specimen upon which this species rests I would designate the
specimen shown on Plate XXXIII, figs. 2 and 2a.

23. Vaccinium pateocorymbosum Jennings, sp. nov. (Plate XXXIII, Figs.

3 and 3a, Type, and 4, 4a.)

The type specimen consists of an impression of a leaf with the apex broken off.
The petiole is about 0.7 mm. in diameter, 4 mm. long, curved, uniform throughout
its length. The leaf-blade is ovate-lanceolate, widest at about the middle, where it is
1.8 mm. wide, apparently about 4.5-5 cm. long, rather bluntly tapering towards
the petiole, somewhat more narrowly tapering toward the apex; the margins
entire, slightly revolute. The midrib is rather strong, tapering above, the secon-
dary veins pinnate in probably about eight approximate pairs (six pairs are pre-
served in the specimen), these veins spreading at an angle of about 50° with the
midrib, curving upward towards the margin and finally looping successively with
branches from the lower side of the next vein above. The tertiary venation is
fine and rather indistinct, but toward the margin and along the lower side of the
lowermost pair of veins forming small narrow loops, while between the veins there
are a few tertiaries running across from one vein to the other at almost right angles
to them, the final meshes in the interior of the blade being rather small and indis-
tinct. There appears to be a very narrow decurrent flange along each side of the
petiole tliroughout its length.

In size, shape, and venation the specimen almost exactly matches leaves
which can be selected from the modern Vaccinum corymbosum Linn^us of swamps
and low woods from Maine, Quebec, and Minnesota to Virginia and Louisiana,
about the only noticeable difference being the somewhat larger angle of divergence
of the veins in the fossil specimen. In length of petiole, number and spacing of the
veins, narrow loops on the lower side of the lowest pair of veins, revolute margin,
margined petiole, etc., the resemblance is too close to be ignored. At first glance
the fossil suggests Quercus cinereoides Lesquereux^® but in that species the base is
more rounded, the venation does not match that of the Missoula specimen very
closely, and there is little basis for believing that an oak would have such a margined
petiole.

Lesquereux, Leo. The Tertiary Flora, U. S. Geological Survey of the Territories, VII, 1878,
p. 152, PL 21, fig. 6.

JENNINGS: FOSSIL PLANTS FROM VOLCANIC ASH BEDS, WESTERN MONTANA. 427

The genus Vaccinium now contains about 130 species and it ranges from the
Arctic Circle to the mountains of the tropics, having its greatest development in
the Himalaya Mountains and in North America. Considerable doubt has been
expressed as to the correct identification of some of the fossil forms reported, but
it seems fairly certain that the genus is correctly ascribed to the Eocene, with three
or four species from Alaska and Colorado, while in western Europe in the Oligocene
and, especially, in the Miocene, the genus was represented by a number of species,
so that the occurrence of Vaccinium in the Missoula region in the Oligocene would
not be unexpected.

428

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXII.

Fig. 1. Buds of Equisetum sp. Enlarged two and one-half diameters.

Fig. 2. Equisetum insculptum Jennings. Type. Enlarged two and one-half
diameters.

Fig. 3. Sequoia Haydenii (Lesquereux) Cockerell. Natural size.

Fig. 3a. The upper part of the same specimen, enlarged two and one-half diameters.

Fig. 4. Sequoia Haydenii (Lesquereux) Cockerell. Enlarged about two diameters.

Fig. 5. Sequoia Haydenii (Lesquereux) Cockerell. A young shoot enlarged
three and one-half diameters.

All photographed from specimens in the Carnegie Museum collected by Earl Doug-
lass from Montana. Figs. 1 and 2 from “Locality 139,” Winston; Figs. 3-5 from
“Locality 196,” near Missoula, Montana.

Memoirs Carnegie Museum, Vol. VIII.

Plate XXII.

Equisetum and Sequoia.

(For explanation see opposite page.)

430

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXIIL

Fig. 1. Sequoia oblongifoKa Jennings. Type, enlarged about one and three-
fourths diameter.

Fig. la. Part of same branch, enlarged two and one-half diameters.

Fig. 2. Sabina lingucefolia (Lesquereux) Cockerell. Enlarged about two and
one-half diameters.

Fig. 3. Sequoia Haydenii (Lesquereux) Cockerell. Enlarged two and one-half
diameters.

Fig. 4. Sequoia Haydenii (Lesquereux) Cockerell. Cones, enlarged about three
and one-half diameters.

Fig. 5. Thuyopsis gracilis Heer. Enlarged about three and one-half diameters.

Fig. 6. Sequoia Haydenii (Lesquereux) Cockerell. Natural size.

Photographed from specimens in the Carnegie Museum collected by Earl Douglass
from “Locality 196,” Missoula, Montana.

Memoirs Carnegie Museum, Vol, VIII.

Plate XXIII.

Sequoia, Sabina, and Thuyopsis,
(For explanation see opposite page.)

432

MEMOIRS OF THE CARNEGIE MUSEUM,

EXPLANATION OF PLATE XXIV.

Figs. 1, 2, and 3. Sequoia Haydenii (Lesquereux) Cockerell. Different types of
leaves of the same species.

Fig. 4. Betula multinervis Jennings.

Fig, 5. Ilex furcinervis Jennings.

Photographed from a specimen in the Carnegie Museum collected at Missoula,
Montana, from “Locality 196”. About seven-eighths diameter.

Fig. 6. Populus Zaddachi Heer.

Fig. 7. Alnus microdontoides Jennings.

Fig. 8. Alnus Hollandiana Jennings.

Photographed from a specimen in the Carnegie Aluseum, collected at “Locality
196,” Missoula, Montana, by Earl Douglass. About four-fifths diameter.

Memoirs Carnegie Museum, Vol, VIII.

Plate XXIV.

Sequoia, Betula, Ilex, Populus, and Alnus.
(For explanation see opposite page.)

434

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXV.

Fig. 1. Populus Zaddachi Heer. Natural size.

Fig. 2. Populus Zaddachi Heer. About seven-eighths diameter.

Fig. 3. Alnus Hollandiana Jennings. Type. About seven-eighths diameter.
Photographed from specimens in the Carnegie Museum, collected at “Locality
196/’ Missoula, Montana, by Earl Douglass,

Memoirs Carnegie Museum, Vol. VIII. Plate XXV.

Populus and Alniis.

(For explanation see opposite page.)

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436

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXVI.

Fig. 1. Populus Zaddachi Heer. Natural size.

Fig. la. Middle left-hand margin of same specimen, enlarged two and one-half
diameters.

Fig. 2. Populus Zaddachi Heer. Natural size.

Fig. 3, Populus Zaddachi Heer. Base of leaf shown in Plate XXV, Fig. 1, enlarged
two and one-half diameters.

Fig. 4. Populus smilacifolia Newberry. Natural size.

Fig. 5. Populus Zaddachi Heer. Natural size.

Fig. 6. Populus Zaddachi Heer. Another leaf enlarged two and one-half diameters.

All photographed from specimens in the Carnegie Museum collected by Earl Doug-
lass at Missoula, Montana. — Figs. 1, 2, and 3 from “Locality 196;” Figs. 4, 5, and 6
from “Locality 165.”

;• .M

Memoirs Carnegie Museum, Vol. VIII,

Plate XXVI,

Popuhis Zaddachi and P. smilacifoUa.
(For explanation see opposite page.)

438

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXVII.

Fig. 1. Betula multinervis Jennings. About three-fifths diameter.

Fig. la. Same specimen, natural size. Type.

Fig. 15. Part of “la,” enlarged two and one-half diameters.

Fig. Ic. Same as “16,” with the margin of the leaf drawn in.

Fig. 2. Betula multinervis Jennings. Cones, enlarged about two and one-half
diameters.

Fig. 3. Quercus ajrproximata Jennings. Tj^pe. Natural size.

Fig. 3a. Part of the same specimen, enlarged about two and one-half diameters.
All photographed from specimens in the Carnegie Museum collected by Earl Doug-
lass at Alissoula, Montana. Figs. 1 and 3 from “Locality 196;” fig. 2 from “Locality
165.”

Memoirs Carnegie Museum, Vol. VIII,

Plate XXVII.

Betula and Quercus.

(For explanation see opposite page.)

440

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXVIII.

Fig. 1. Alnus Hollandiana Jennings.

Fig. 2. Betula multinervis Jennings,

Fig. 3. Quercus laurosimulans Jennings.

About two-thirds diameter. Photographed from a specimen in the Carnegie
• Museum, collected by Earl Douglass at “Locality 196,” Missoula, Montana.

Memoirs Carnegie Museum, Vol. VIII

Plate XXVIII

Alnus, Betula and Quercus.

(For explanation see opposite page.)

■ ;

442

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXIX.

Fig. 1. Juglans pentagona Jennings. Type. Natural size.

Fig. la. Part of same specimen, enlarged about two and one-half diameters.

Fig. 2. Juglans pentagona Jennings. Natural size.

Fig. 2a. Part of the same specimen, enlarged about two and one-half diameters.
Fig. 3. Cyperacites sp. Enlarged two and one-half diameters.

Fig. 4. Typha Lesquereuxi Cockerell. Enlarged two and one-half diameters.
Photographed from specimens in the Carnegie Museum collected by Earl Douglass
from Missoula, Montana. Figs. 1, la, 2 and 2a from “Locality 165;” figs. 3 and 4
from “Locality 196.”

1

Memoirs Carnegie Museum, Vol. VIII.

Plate XXIX.

Jnglans, Cyperacites, and Typha.
(For explanation see opposite page.)

444

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXX.

Fig. 1. Alnus Hollandiana Jennings. Type. Natural size.

Fig. la. Aliddle right-hand margin of same, enlarged two and one-half diameters.
Fig. 2. Alnus microdontoides Jennings. Type. Natural size.

Fig. 2a. Lower part of same specimen, enlarged two and one-half diameters.

Fig. 3. Alnus cones. Enlarged about two and one-half diameters.

Photographed from specimens in the Carnegie Museum collected at “Locality
196,” Missoula, Montana, by Earl Douglass.

Memoirs Carnegie Museum, Vol. VIII,

Plate XXX,

Alnus H ollandiana and A. ■microdo7it aides.
(For explanation see opposite page.)

446

MEMOIES OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXXL

Fig. 1. Quercus flexuosa 'Newberry. Natural size.

Fig. la. Part of the same specimen, enlarged about two and one-half diameters.

Fig. 2. Celastrus parvifolius Jennings. Type. Enlarged about two and one-half
diameters.

Fig. 2a. Same with the venation inked in.

Fig. 3. Quercus laurosimulans Jennings. Natural size.

Fig. 3a. Part of another leaf from the same place, enlarged about two and one-half
diameters.

All photographed from specimens in the Carnegie Museum collected by Earl Doug-
lass at Missoula, Montana. Fig. 1 from ‘^Locality 165”; figs. 2 and 3 from “Locality 196”.

Memoirs Carnegie Museum, Vol, VIII

Plate XXXI.

Quercus and Celasti'us.

(For explanation see opposite page.)

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448

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXXII.

Fig. 1. Ficus (?) prunifolia Jennings. Type. Natural size.

Fig. la. Part of same specimen, enlarged about two and one-half diameters.

Fig. 2. Ilex furcinervis Jennings. Type. Natural size.

Fig. 2a. Upper part of same specimen, enlarged about two and one-half diameters.
Fig. 26. Same as 2a, but with the margin inked in.

Fig. 3. Acer oregonianum Knowlton. Enlarged about two and one-half diameters.
Photographed from specimens in the Carnegie Museum collected at Missoula,
Montana, by Earl Douglass; figs. 1 and 2 from “ Locality 196 ”, fig. 3 from “Locality 165 ”.

Memoirs Carnegie Museum, Vol, VIII,

Plate XXXII.

Ficus, Ilex, and Acer.

(For explanation see opposite page.)

450

MEMOIRS OF THE CARNEGIE MUSEUM,

EXPLANATION OF PLATE XXXIII.

Fig. 1. Aralia longipetiolata Jennings. Natural size.

Fig. la. Same, but with the venation inked in.

Fig. 2. Aralia longipetiolata Jennings. Base of a larger leaf, natural size. Type.
Fig. 2a. Same, with the venation inked in.

Fig. 3. Vacciniurn palceocorymhosum Jennings. Type. Natural size.

Fig. 3a. Same, enlarged about two and one-half diameters.

Fig. 4. Vacciniuin palceocorymhosum Jennings. Natural size.

Fig. 4a. Same, enlarged about two and one-half diameters.

All photographed from specimens in the Carnegie Museum, collected by Earl
Douglass. Figs. 1 and 2 from “Locality 139”, Winston, Montana; figs. 3 and 4 from
“ Locality 196 ”, Missoula, Alontana.

Memoirs Carnegie Museum, Vol, VIII,

Plate XXXIII.

Aralia and V accinium.

(For explanation see opposite page.)

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44. Additions to Flora of Allegheny County, Pa.

Jennings ................................ .15

45. New Species of Kneiffia. Jennings .......... .05

46; Occurrence of Triglochln palastris m Pa. Jen-
nings .................................... .05

47. New Species of ' Ibidiam. Jennings ......... .10

48. Agate Spring Fossil Quarry. Peteeson ...... .10

49. Two MTew Birds from British E. Africa. Obee-

HOLSEE ................................... .05

50. The Ohazy Formation and Its Fauna. Eay-

' mond .................................... 1.60

51. A New American Cybele. Naseawat and Eat-

mond .................................... .16

52. Plastron of the ProtosteglnBS. Wieland ..... .15

53. New Species of Testudo from the Miocene, etc.

0. P. Hat- ................................. .25

54. Miocene Beds of W. Nebraska and B. Wyoming

and Their Vertebrate Faun®. Peteeson .... 1.00

55. New Species of Lonicera from Pa. Jennings. .05

56. Merycochoerus, etc. Douglass.

57. New Meryeoidodonts. Douglass, (Nos. 56

and 57 sold together.) ..................... 1.00

58. Further Collections of Fishes from Paraguay.-

■ Eigenmann, McAtee, and Waeb ............ 1.26

69. Undetermined Element in the Osteology of the

Mosasaurld®. Holland ............. ,20

€0. Gasteropoda of the Ohazy. Eaymond ........ 1.35

61. Occurrence of V/ymea Americana in Pa. Jen-

nings .................................... .05

62. Account of Pleistocene Fauna Discovered at

Frankstown, Pa. Holland ................ .10

63. Vertebrate Fossils from the Vicinity of Pitts-

burgh, Pa. Case ......................... .15

64. lU^nid® from the Black Eiver Limestone near

Ottawa, Canada. Eaymond and Nabeawat. . .20

65. Ehinoceroses from the Ollgocene and Miocene

of North Dakota and Montana. Douglass . . .26

66. Fossil Horses from North Dakota. Douglass. .30

67. Oligocene Lizards. Douglass ............... .20

68. The Type of Stenomylus gracilis. Peteeson. . .25

69. New Species of Birds from Costa Eiea, etc.

Caeeiker ................................. .10

70; Costa Elcan Fonnlcarlid®. Carriker ........ .05

71. Vertebrate Fossils from the Fort Union Beds.

Douglass ................................ .45

■’ of the LepMoptera of Western Pa., etc.

. GEL ................................... 1.25

la of Upper Devonian in Montana. Fossils
Bed Shales. Eaymond ................. .60

■ ■ Species of Procamelus from Upper Mio-
- ■ e of Montana. Douglass ............... .30

.of the Conemaugh Series between
Pittsburgh and Latrobe, Pa. Eaymond ..... .36

76. List of the Unionld,® of Western Pa. Ortmann ,50

77. Geological Reconnaissance in North Dakota,

Montana, and Idaho. Douglass ........... 1.00

78. Botanical Survey of Presque Isle. Jennings . . 2.75

79. Sesqul-Centennial (Pittsburgh) EeHcs. Stewart .46

80. Dromomeryx, New Genus of American Rumi-

nants, Douglass ......................... .30

.. 4 . : n. • 'I eial Drift and Devonian and

' ■ jar Meadville, Pa. Millard. . .10

- Helodus, Eastman .......... .05

’'V . -'--:'..; y Mellor. Holland.......... .15

r - j- , sdltion to British Guiana, etc.

' ■ • ' :■■■■-■ ............................. .60

85. Reports of Expedition to British Guiana, etc.

Durbin .25

86. Odonata of the Neotropical Region, Exclusive

of Mexico and Central America. Calvert . . . 2.25

87. Deinosuchus hatcheri, a New Genus and Species

of Crocodile. Holland ................... .20

88. Reports on Expedition to British Guiana, etc.

Blosseb ................................ . .15

89. Description of Some New Titanotheres from

the Uinta. Douglass ..................... ,25

90. Annotated List of the Birds of Costa Rica In-

cluding Cocos Island. Carriker 3.00

91. Geology of the Coast of the State of Alagoas,

Brazil. Beanner ......................... .40

92. Fossil Fishes from the Bituminous Shales at Ei-

acho Doce, Alagoas, Brazil. Jordan ........ .55

93. Ordovician Trilobites, No. H. Asaphid® from

the Beekmantown.- Raymond .............. .35

94. Ordovician Trilobites, No. III. Raymond &

Nareaway ............................... .85

95. Ordovician Trilobites, No. IV. Raymond .... ,40

96. Fishes from Irkutsk, Siberia. Jordan and

- Thompson ............................... ■ .30

97. South American Tetrigid*. Bruner ......... 1.00

98. Fauna of the Allegheny and Conemaugh Series

in Western Pa. Raymond ................. .30

99. Ichthyological Survey About the San Juan

Islands, Washington. Starks ............. .75

100. New Species of Pygldlum. Eigenmann ...... .10

101. The BxacMopoda and Ostracoda of the Chazy.

Raymond .50

102. A New Camel from the Miocene of Western

■ Nebraska. Peterson ..................... .15

103. Skeleton of Stenomylus Mtchcoeki, etc. Petee-

son ...................................... .15

104. Skeleton of Diceratherium cooki. Peterson . . .16

105. Carnegie Museum Expedition to Central South

America, 1907-1910. Holland ............. .15

106. Report Upon the Expedition of the Carnegie ■

Museum to Central South America. Hase-
MAN & Eigenmann ....................... .25

107. New Species of Fishes, etc., from Central South

America. IIaseman ...................... .50

108. Annotated Catalog of Oichlld Fishes from Cen-

tral South America. Haseman ............ 1.25

109. New SpQcies of Pishes from the Rio Iguassu.

Haseman ................................ .65

110. Ornithology of the Bahama Islands. Todd &

Worthington ............................. .76

112. South American Acrldoidea. I. Bsunek .... 1,00

113. Species of Hasemaaia, Hyphessobrycon, and

Hemigrammus. Ellis ..................... ,25

114. New Ohaxacins in the Carnegie Museum. Eigen-

MANN ..................................... .30

115. Jurassic Saurian Remains Ingested within Fish.

Eastman ................................ .20

116. Autograph Letter of U. S. Grant to Edwin M,

Stanton. Holland ........................ .16

117. Albert J. Barr. By W. J. Holland ........... .10

118. Seventeen New Neotropical Birds. Todd ..... .26

119. Dr, David, Alter, the First Discoverer of Spec-

trum Analysis. Holland .................. .10

120. Two Mummy Labels. Allen ............... .10

121. The Families and Genera of Najades. Ort-

mann ..................................... 1.00

122. Group of StenomyHns in the Carnegie Museum.

Peteeson .26

123. Tertiary Fisli-remains from Spanish Guinea,

Eastman .25

124. Plated Nematognaths. Ellis 65

125. New Species of Camharus from the Isle of

Pines. Ortmann .16

126. Sedum Carneglei, a New Species of the Family

Crassulacese, etc. Hamet 10

127. Two New Species of Fishes from Peru. Eigen-

MANN 15

128. South American Locusts (Acridoidea). 11.

Bruner 75

129. Revision of the Genus Chsemepelia. Todd 85

130. New Genus and Species of Abyssinian Rodents.

Childs Frick 20

131. New Titanothere from the Uinta. Peterson.. .20

132. Small Titanothere from the Lower Uinta. Peter-

son .10

133. Osteology of Lasiopyga and Callithrix, etc.

Shupeldt .26

134. New Ehynchocephalian from Solenhofen. Grier .15

135. Scales of South American Characinid Fishes.

Cockerell 25

136. Skeleton of Platigonus Leptorhinus. Peter-

son 10 .

137. Lichens Collected in the Thunder Bay District,

Ontario. R. Heber Howe, Jr .10

138. Preliminary List of Fossil Plants in the Roof

of the Pittsburgh Coal. Grier ,10

139. Undescribed Remains of the Uinta Titanothere

Dolichorhinus. Peterson 15

140. Triassic Fishes Belonging to the Catopteridse

and Semionotidffl. Eastman .15

141. Osteology of Promerycochoeims. Peterson ... .40

142. Correction of a Generic Name. Peterson 05

143. Skull of Bison Crassicornis. Holland 10

144. Serrasalminse and Mylinie, Eigenmann .50

145. Heads and Tails: Notes on Sauropod Dinosaurs.

Holland 15

146. Dipterus Remains from Upper Devonian of

Colorado. Eastman 10

147. Notes on Tropical American Tettigonoidea (Lo-

custodea). Bruner 1.65

148. New Species of Tortoise from Jurassic of

Utah. Gilmore 16

149. Fauna of Upper Devonian in Montana.

Haynes .45

150. New Sphagebranchus from Bahamas. Eigen-

mann 07

151. Marine Fishes from Colombia and Ecuador.

Wilson 16

152. Apareiodon, a New Genus of Characid Fishes.

Eigenmann 18

153. New and Rare Fishes from S. American Rivers.

Eigenmann 25

154. Three New Species of Characid Fishes. Eigen-

mann & Henn 12

155. The Species of Salminus, Eigenmann 7

156. South American Poeciliid Fishes. Henn 40

157. A New Species of Apatosaurus. Holland .... .7

158. Birds of the Isle of Pines. Todd ............ ,70

159. Reptiles and Amphibians of the Isle of Pines.

Barbour 20

160. Land and Fresh Water Shells of the Isle of

Pines. Henderson 20

161. Pelecypoda of the Chazy. Raymond 45

1 62. S. American Crickets, Gryllotalpoidea, and Ache-

toidea, Bruner 1.30

163. Preliminary Catalog of N. American Sphsen-

idse. Sterki 75

164. Directions for Collecting Sphseriidae and

Aquatic Gastropods. Sterki ... ,10

165. Lepidoptera of the Isle cf Pines. Holland. ... .60

166. Odonata of the Isle of Pines. Kahl 15

167. A Trip to Islands in Lake Erie. Goodrich 15

168. Land-Shells of Islands at Western End of Lake

Erie, etc. Clapp 20

169. Orthoptera of the Isle of Pines. Holland &

Kahl 16

170. Obituary Notes. G. A. Link, T. A, Mills, Boyd

Crumrine, E. M. Bigelow. Holland 10

171. Two New W. African Rhopalocera. Holland. $0.15

172. Botany of the Isle. of . Pines. Jennings 2.00

173. List of Hymenoptera of Isle of Pines. Holland .10

174. Some Species of Farlowella. Eigenmann .... .30

175. List of S. American Lizards. Griffin ...... .35

176. Leptodeira albofusca (Lacepdde). Griffin.... ,10

177. List of Coleoptera collected on Isle of Pines.

Holland & Schwarz 15

178. Rhjmchota of the Isle of Pines. Heidemann

& Osborn ’ "

179. Mammals of the Isle of Pines. Holland .... .(

180. Report upon FossR Material Collected in Cave

in the Isle of Pines. Peterson

181. New Species of Fern (Polystichum jenningsi).

Hopkins ]

182. Notes upon the Genus Leucophenga Mik, with

Descriptions of New Species. Kahl 30

183. Some Species of Rhamdia, a Genus of S,

American Siluridas. Eigenmann & Fisher. .15

184. New and Rare Species of. South American

Siluridse. Eigenmann 30

18'5. List of the Hypopthalmidse, Diplomystidse, and

some Unrecorded Species of SUuridse. Fisher. .30

186. A Synopsis of the Saurian Genus Prionodac-

tylus. Griffin ' 05

187. Notes on a Collection of Fishes from Ceylon

with Descriptions of New Species. Jordan
& Starks •' .35

188. Collection of Fishes from Port Said, Egypt.

Jordan & Hubbs 20

189. A Fossil-bearing Alluvial Deposit in Saltville

Valley, Virginia. Peterson

190. Catalog of Collection of Watches belonging

to H. J. Heinz. Stewart, Holland, Oogges-
hall Paper, BO cts.; hoards

191. Contributions to the Natural History of the

Isle of Pines: Reprinted from Annals, Vols.
VIII-XI, 560 pp., 34 plates, index. Bound
in cloth 5.00

192. Material Discovered in the Upper Eocene of

the Uinta Basin by Earl Douglass and O. A.
Peterson. Peterson 2.25

193. The Fresh-Water Fishes of the Island of For-

mosa. Masamitsu Oshima 2.75

194. On Elephenor, a New Genus of Fishes from

Japan. David Starr Jordan

195. A Description of CypripediU'

Jennings

196. Obituary of Charles Rochester

197. Obituary of Andrew Arnold La

198. Obituary of Herbert Huntin^oi

199. Obituary of Henry John Heinz

Publications of the Carnegie Museum, Serial No. 109

MEMOIES

OF THE

CAENEGIB MUSEUM

VOL. Yin

W. J. HOLLAND, Editor

No. 3

// \ \
(* NOV 2 6 1921 ' *!

■%'onal

SOUTH AMERICAN NAIADES; A CONTRIBUTION TO THE
KNOWLEDGE OP THE FRESHWATER MUSSELS
OF SOUTH AMERICA

BV

DR. A. E. ORTMANN

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MEMOIRS

. OF THE

CARNEGIE MUSEUM

VoL. VIII. No. 3.

SOUTH AMERICAN NAIADES; A CONTRIBUTION TO THE
KNOWLEDGE OF THE FRESHWATER MUSSELS
OF SOUTH AMERICA.

By Dr. A. E. Ortmann.

(Plates XXXIV-XLVIII)

Introductory.

During the expedition of the Carnegie Vluseiim to central South America,
from 1907 to 1909, Mr. J. D. Haseman had as his prime object the collection of
Fishes (Haseman, 1911)h At the request of the present writer he, however, took
particular pains to collect and preserve freshwater mussels. The result was one
of the finest and largest collections of South American Naiades ever secured. The
value of this collection is enhanced by the fact that a great number of s})ecimens
were preserved in alcohol with the soft parts, and the study of their anatomy has
thus been made possible. Preliminary notes concerning the most important points
of structure have been previously published (Ortmann, 1911a), chiefly in order to
set forth the affinities of the South American forms with those of the rest of the
world. Only a few typical forms were selected and discussed for this purpose.
Further examination of the material has revealed a number of additional and highly
interesting facts with regard to the anatomy, which throw light on the taxonomy
and phylogeny of this group.

' The references in parentheses are to the papers found in the biblioo'raphj^ at the end of this paper.

451

452

MEMOIRS OF THE CARNEGIE MUSEUM.

The present paper is intended to give a full account of these investigations.
Naturally all accessible material has been included. In many cases only shells
without their soft jiarts were available. This remark refers largely to older and
some more recent material preserved in the Carnegie Museum obtained from various
sources. The Haseman Collection likewise contains only the hard parts of many
species, but on the other hand, other accessions consisted of shells with soft parts,
as, for instance, the siiecimens of Ajiodontites crispaius from the upper Magdalena-
drainage in Colombia, collected by Dr. C. H. Eigenmann, and a collection of a
number of species from the La Plata in Argentina, received from Dr. A. Wind-
hausen.

In spite of the comparative wealth of material at hand, in some respects our
collections are not complete and do not permit the determination of certain ques-
tions, chiefly taxonomic. My greatest difficulty has been to properly identify
the species” at hand. The older writers, Spix, Wagner, Lea, D’Orbigny, Philippi,
Hupe, and others, generally described their “species” from very insufficient ma-
terial, and the author to whom we are most indebted for clearing up the taxonomy
of the South American Naiades, H. Von Ihering, also was often handicapped by
having too scanty material. All previous writers had no clear conception of the
range of variation in the various forms, and thus their descrijitions generally are
those of individuals, often indeed very elaborate and complete, but without proper
emphasis laid upon the really important specific characters. Simpson’s “Descrip-
tive Catalogue” (1914) did not much improve matters, since he was largely de-
pendent iqion the unsatisfactory publications of previous authors.

I do not claim, by any means, that my treatment of these shells overcomes all
these difficulties; on the contrary, I am in many particulars not at all satisfied
with the results obtained. Nevertheless, I claim that the study of the soft parts
of a great many forms has furnished a basis for the proper understanding of what
the “species” are, and has, at least, furnished a clue to their systematic arrange-
ment, incom])lete and fragmentary, it may be, but which probably wall prove to
be of great value, when the soft parts of all or most of the species are known.

The difficulty encountered in recognizing the described forms is sometimes
exas]ierating. We should expect, in cases where exact type-localities are given,
and where material from these is at hand, that it would not be very great. But,
for instance, even of the species described by Von Ihering from Sao Paulo, I have
recognized only a com]iaratively small number, although I ])ossess a large quantity
of material rejn-esenting the Naiades from that state. In other cases, when the
t3'pe-locality is vaguel^q or not at all, given, it has been practically impossible to

ortmann: south American naiades.

453

be sure of the identification. In consequence I have been compelled to introduce a
number of “new species,” although I am afraid that some of them are not really
“new.” But I must leave the task of making out their synonymy to others, who
have access to authentic material representing the older forms. It should also
be remembered that the introduction of new names is justified by the rules of
nomenclature, when an original description is insufficient to enable the species to
be recognized .

General Remarks as to the Affinities and Geographical Distribution of

THE South American Naiades.

The earlier writers generally placed the South American forms in the old
collective genera, Unio and Anodonta, to which a kind of intergrading group, called
Monocondylcea, and certain specialized types, such as Hyria and Castalia, were
added. Von Ihering was the first to recognize that the South American “Ano-
donta” so-called, differs from the Anodonta of the northern hemisphere in important
characters, and that it is related to certain African forms, Alutela, Spatha, etc.
He calls this genus Glabaris Gray = Anodontites Bruguiere. But he left the other
forms under Unto. For these Simpson (1900) used the name Diplodon Spix. In
my preliminary report on South American Naiades (Ortmann, 1911a, pp. 108, 120,
129, 130) I was able to show that Diplodon, as well as Hyria and Castalia, differ
anatomically from the Unionidm of the northern hemisphere, and that Von Ihering’s
separation of Glabaris from Anodonta is fully justified and correct. I also found
that the genera Hyria, Castalia (= Tetraplodon), and Diplodon, recognized by
Simpson as a peculiar group, but still placed with the Unionidce, are actually more
nearly allied to “Glabaris,” and form with this a group, the family Mutelidce, which
should be divided into two subfamilies, the Hyriinoe and Mutelince, each with a
number of genera, the anatomy of many of which, however, was still unknown.

Subsequent investigations brought out the fact that Simpson (1900) was cor-
rect in associating certain Australian Naiades with the South American Diplodon,
since I was able to show that D. australis has practically the same anatomical
structure as the South American Diplodon (Ortmann, 1912), but that it should be
elevated to the rank of a separate genus, Hyri della Swainson, admitted as a sub-
genus by Simpson.

The systematic arrangement and geographical distribution of the families and
subfamilies of the Naiades would thus be as follows:

454

MEMOIRS OF THE CARNEGIE MUSEUM.

Superfamily NAIADES.

I. Family Margaiutanid/e Eurasia and North America (discontinuous).

TI. Family Unionid.®:

1. Subfamily Unionin.e Eurasia, Africa, North America, southward to Central America.

2. Subfamily Anouontin.e Eurasia, North America, southward to Central America.

3. Subfamily Lampsilin.e North America southward to Mexico.

III. Family Mutelid.e;

1. Subfamily IIyriin.e South America (but not in Central America), Australia.

2. Subfamily Mutelin.e South America to southern Mexico, Africa.

I can not imjirove u]ion this arrangement at present. It possibly might be
advisable, in future, to elevate the two South American subfamilies to the rank of
families, but I refrain from doing this, chiefly because the African and Australian
forms belonging to them are too ])oorly known. The fact that in South America
two grouiis of Naiades are found, which are more closely allied to each other than
to any other groiij), is well exju-essed by uniting them into one family (Midelidce) .

The more iirimitive subfamily, Hyriince, is found all over South America, but
becomes rare in the northern jiarts (Venezuela and Colombia), and is missing, so
far as known, in Central America. It is (piite almndant in Chile, where the Mu-
telirm are absent, and it is also found in Australia.^

The more specialized group, subfamily Mutelince, is found in South America,
east of the Andes. It is missing in Chile, but has been rejiorted from the Pacific
drainage in Ecuador. It goes into Central America and reaches southern Mexico.
On the other hand it is represented in Africa, probably all over the continent,
with the excejition of the Mediterranean region.

This geograiihical distribution of the larger groups is extremely significant,
for it serves to support certain theories as to the origin of the South American
continent, its former connections, and the origin of its fauna. The presence of
the Hyriince both in South America and Australia indicates the former connection
of both continents, probalily by way of Antarctica, and the fact that species of
Diplodon, the most primitive of the South American Hyriince, are found in Chile,
is entirely in keeping with this. The fact that Uriionidce with certain Hyriine
structures^ are found chiefly in southeastern Asia, suggests that they probably

^ It is unknown whether all the Australian Naiades belong here. Unionidce related to the Hyriince
in having the septa of the marsupiuin interrupted, are known to me from southeastern Asia (Siam,
China) and northwestern America (Pacific slope), but these forms certainly are Unionidce in all other
resjjects.

® Interrupted septa in the marsupium are known in the Asiatic Unionmce: Lamellidens (Ortmann,
1911a, p. lOb); in Hyriopsis and Contradens (Ortmann, Nautilus, XXX, 191G, i)p. 85 and lOG); and,
according to the figures of Haas (System. Conchyl. Cabin., XIX, 1911 and 1912), also in Rectidens and
Acuticostn.

ortmann: south American naiades.

455

originated on the old connecting land between Asia and Australia (Sino-Aiistralian
continent) or in Australia, and that Diplodon first reached Chile (in Mesozoic times)
coming from Australia, and subsequently invaded the Brazilian mass. The
modern forms of the Hyriince {Castalia, Hyria) chiefly have their center in the
basin of the Amazon, a comparatively young part of the continent.

The more advanced type, represented by the subfamily Mutelinm, developed
probably in Brazil at the end of the Mesozoic, when it had a chance to spread
over the old connection across the Atlantic (Archhelenis of Von Ihering), and the
immigration of this stock into Central America is of rather late date (late Tertiary).
It is not very likely that the Mutelinm reached South America coming from Africa,
because there is no trace of them found in southern Asia.'^

Diagnostic Characters of the South American Naiades.

(See Text-figures 1, 2, 3.)

Family: MUTELID.Tl Ortmann (1911).^

1. Diaphragm between branchial and cloacal cavity formed anteriorly by the
gills, posteriorly by a solid union of the mantle margins (Figs. 1, 2, 3, t). (No such
mantle connection in Unionidoc).

2. Anterior end of inner gill (Figs. 1, 2,

3, i) broadly attached, and in contact with
posterior base of palpi (h). (In the Union-
id(B, there is always a longer or shorter space
between these parts.)

3. Anal and branchial openings (a and
b) sharply separated from each other by the
union of the mantle margins (See Character
1); anal opening open or closed above, but
there never is a supra-anal opening. (In the
Unionidce, the anal may be open, but, when
closed, there is always a supra-anal opening.)

■* In Africa there are also Unionidce of the subfamily Unionince; they undoubtedly point to a con-
nection with Asia, since such forms are plentiful there, and this indicates a different route of immigra-
tion from that of the Mutelince.

The name depends upon the investigation of the anatomy of the African genus Mulela, which is
unknown, but we have every reason to assume, chiefly through Von Ihering’s study of the shell, that
this genus will fall under this family. The family name Miitelidce was first used by II. & A. Adams
(1858), but for an entirely different association of forms. The Mutelidce of Simpson (1900) correspond
to our Mutelince.

Fig. 1. Diagram of soft parts of female
of Diplodon trijidus (Lea). Natural size,
left section of mantle removed, a, Anal
opening; 6, Branchial opening; h, Palpi; i,
Inner gill; o. Outer gill; p, Pes; s. Closed
l)art of anal opening; t, Union of mantle
separating anal and branchial openings.

456

MEMOIRS OF THE CARNEGIE MUSEUM.

4. Gills with or without water-tubes, and with isolated, scattered inter-
laminar connections, or with interrupted or solid septa. (In the Unionid<je, mostly
uninterrupted sejita and water-tubes are found; very rarely are they interrupted;
the forms with the latter jirobably form the connection with the Mutelidce.)

5. Marsupium only in the inner gill (Figs. 1, 2, 3, i). (In the Unionidce, the
marsupium is either in all four gills, or in the outer gills, never in the inner gills
alone.)

G. Certain advanced genera of the Mutelidce show a tendency to close the
branchial opening in front by a mantle connection. (Such a connection is entirely
unknown in the Unionidce.)

The foregoing are the anatomical characters. The shape of the shell and its
parts are subject to so many variations, both in the Mutelidce and the Unionidce,
that it is practically imiiossilile to point out general differentiating characters.
Different and often very peculiar types of shell, occurring independently, are fre-
quently observed, so that it is clear that the shells by responding to certain stimuli

and requirements have acquired sim-
ilar shapes in forms, which belong to
different families (convergency or
parallelism). Simpson (1900 and
1914) found it impossible to give
shell-characters for his subfamily
' ‘ Hyriance ’ ’ = Hijriince + certain Uni-
onince, except beak-sculpture, and he
was mistaken in this. In our arrange-
ment, this is the only character,
which might be mentioned, but not
without qualification. We may ex-
press it thus: The Mutelidce have ra-
dial beak-sculpture, if such is present
at all);*’ while the Unionidce rarely
have distinct radial beak-sculpture,
but commonly other types, zig-zag, double-looped, or concentric.

Some Mutelidw have, indeed, . shells which completely mimic those of certain
Unionidce. Species of Diplodon often externally so much resemble certain species

® But even this should be (jualified. In certain species of A nodontiles in the subfamily Mutelince
I have observed sometliing like concentric beak-sculpture, while, as a rule, the Mutelmce have no beak-
sculpture whatever.

Fig. 2. Diagram of the soft parts of female of
Castalina nehringi Von Jhering. Natural Size, left
section of mantle removed. (Lettering- as in Text-
figure 1, p. 4559

ortmann: south American naiades.

457

of the North American Elliptio, that without a minute examination of muscular
impressions, hinge-teeth, etc., it is impossible to recognize them, when the locality
is unknown. The only reliable character in this case, beak-sculpture, is often
obliterated by erosion. Species of Anodontites often look like species of Anodonta.
Here the examination of beak-sculpture, muscular impressions, and the ligamentinal
sinus establishes their affinity. On the other hand, there are MidelidoB which are
easily recognized by the peculiar shape of the shell, and could never be confounded
with UnionidcB {Castalia, Hyria, Mycetopoda, etc.). .

The characters of the two subfamilies of the Mutelidm are the following:

Subfamily Hyriin^ Ortmann (1911).

Hyriance Swainson (1840), very closely corresponds to this. Lamphoramphus-

group of Hyriance Simpson (1900).

1. Anal opening (Figs. 1, 2, a) closed above (s), slit-like (in South American
forms), or forming a short, tubular siphon (Australian forms).

2. Marsupium generally an interrupted network of interlaminar connections
(Figs. 1, 2, i), the connections often standing in rows, thus forming incomplete
septa and incomplete, communicating, water-tubes. In rare cases, the inter-
laminar connections of the marsupium (but not of the non-marsupial gills) form
solid septa and isolated water-tubes. Marsupial part of gill often restricted to
only a section of it.

3. Non-marsupial gills always with poorly developed, scattered interlaminar
connections. (Figs. 1, 2, o.)

4. Inner lamina of inner gills always entirely connected with abdominal sac.

5. Palpi (Figs. I, 2, h) subtriangular or subfalciform, with gently curved lower
margins, and somewhat produced posterior points.

6. Larva a glochidium (See Fig. 4, p. 469).

As a character of the shell may be mentioned the beak-sculpture, which is
generally present (often poorly develojied, rarely absent), and always radial.
The hinge-teeth are always present, and generally well-developed. Dorsal muscle-
scars are present.

Subfamily Mutelin^ Ortmann (1911).

MutelidcE Adams (1858), Simpson (1900). The name depends, of course, on the

genus Mutela, the anatomy of the soft parts of which, as has been pointed out

already, remains to be investigated.

1. Anal opening (Fig. 3, a) open (in South American forms, except Myceto-
poda) or closed above (in African forms and Alycetopoda) , slit-like.

458

MEMOIRS OF THE CARNEGIE MUSEUM.

2. Marsiii)iiira (Fig. 3, ^) with well-developed, continuous septa, forming well-
defined, isolated water-tubes, with a peculiar longitudinal ridge or swelling on the
septa near the outer lamina of the gill (PI. XLVIII, figs. 6, 8). When gravid,
only the inner compartment of the water-tubes (towards the inner lamina) is some-
what extended and filled with eggs; the outer compartment becomes a secondary
water-tube (PI. XLVJII, fig. 7h).

3. Non-marsupial gills (Fig. 3, o) also with septa and water-tubes, but the
sejita less strongly developed and without a ridge.

4. Inner lamina of inner gills entirely
connected with abdominal sac (South Ameri-
can forms), or free from it (African forms).

5. Palpi nearly semicircular, longer than
high, with a short posterior truncation,
strongly curved lower margins, and indis-
tinct posterior points (Fig. 3, h).

G. I^arva supposed to be a lasidium
(according to Von Ihering).

The shell generally has no beak-sculp-
ture whatever. In very rare cases a trace
of concentric sculpture has been observed
(see under Anodontites trapezea). Hinge-
teeth more or less obsolete, reduced in num-
ber, or size, and very often entirely absent (Anodontine type of hinge). Dorsal
muscle-scars mostly absent in South American forms, very rarely a faint trace
seen, or a few are present, as in Leila. In African forms, there is one single well-
developed dorsal muscle-scar.

Subfamily HYRIIN.E Ortmann.

^ General Remarks.

Shell of various shajies, subelliptical, subtrapezoidal, subovate, suborbicular,
or sub triangular, sometimes more or less alate. Beak-sculpture mostly present,
rarely missing, but often indistinct or obliterated liy erosion, always of the radial
type, with two sets of radial ridges, starting from two points, immediately in front
and immediately behind the tip of the umlio, and extending to a varying degree
upon the disk. The posterior ridges of the anterior set, and the anterior ridges of
the posterior set, generally interfere with each other in the middle of the disk.

Fig. 3. Diagram of soft parts of female of
Anodo7)tites palagonica ruhicunda (Lea). Nat-
ural size, left section of mantle removed, a,
Anal oi)ening; b, Branchial opening; h, rali)i;
i, Inner gill; o, Outer gill; p, Pes; t, Union of
mantle separating anal and branchial openings.

ortmann: south American naiades.

459

coming there into contact in a sharp angle. There are sometimes irregularities
in the beak-sculpture obscuring their radial character.

Hinge-teeth always present, but rather variable in number, size, and shape.
Normally, there are, as in most other Naiades, two laterals in the left valve, and
one lateral in the right valve, but individual variations may occur in this respect.
The pseudo-cardinals are much more variable. Originally their arrangement
seems to be similar to that found in primitive Unionidce: that is to say, there are
two in the left valve, and one or three in the right; of the latter, the middle one
may be the main tooth, fitting in between the two teeth of the left valve, and the
anterior and posterior tooth may be accessoiy. However, this arrangement is
very often changed by the suppression of certain teeth, and the addition of others.
The most general condition is when of the two pseudocardinals of the left valve
the posterior is more or less obsolete, and often entirely wanting, and only the
two anterior teeth of the right valve are present. Thus the left valve appears
to have only one pseudocardinal {Aspidon of Von Ihering) and two laterals, and
the right valve has two pseudocardinals {Dexion and Epidexion) , and one lateral.
This is chiefly the case in certain species of Diplodon, where the pseudocardinals
are greatly compressed, and directed more or less parallel to the hinge-line, and
this should be kept in mind for the distinction of those species of Diplodon, which
resemble in shape certain North American species of Elliptio, where the normal
arrangement shows two pseudocardinals in the left, and one in the right valve.

This is not the only variation. Additional teeth may turn up, and chiefly when
in the left valve anterior to (or above) the anterior pseudocardinal an additional
tooth is developed {Epaspidon, Von Ihering).

The names Aspidon and Epaspidon, Dexion and Epidexion, may be used to
advantage; but we should not forget that originally there is in the left valve a
tooth behind the Aspidon (regarded by Von Ihering as being accessory), and that
there is often in the right valve another (third) posterior tooth, behind the Dexiond

Further, there is much variability in the shape of the teeth. The pseudo-
cardinals, as has been stated, are often compressed, lamellar, and smooth. But
often they are more solid, or stumpy, when they are frequently more or less split
and divided, sometimes almost cut up into a number of teeth. The laterals may
be smooth or corrugated, the corrugations standing obliquely on the faces; and in
certain genera there are characteristic regular and parallel striations or ridges

^Compare also Odhner, 1918, p. 574 ff. (Homologies of hinge-teeth of “ Unioiiidw”). Odhiier
seems to be right in a general way, but the hinge of the Naiades is extremely complex and individually
variable.

460

MEMOIRS OP THE CARNEGIE MUSEUM.

standing vertically to the edge of the teeth. Similar ridges may be present on
the pseiidocardinals.

In the shape of the muscle-scars on the inside of the shell, there exists no
great difference between the Hyrimce and Mutelincc. The general rule is in the
Hyrimte that the upper anterior retractor-scar is distinctly separated from the ante-
rior adductor impression, while the lower anterior retractor-scar is confluent with the
latter. Very rarely the lower anterior retractor-scar is isolated. (In the Mutelince
either all three scars are confluent, or the lower anterior retractor-scar is isolated;
the upper anterior retractor-scar is hardly ever isolated) . The posterior muscle-scars
are confluent, the posterior retractor-scar forming a small process at the upper margin
of the adductor-scar. The muscle-scars may be deeper or shallower, the upper an-
terior retractor-scar generally is rounded, small, and remarkably deep.

Dorsal muscle-scars are present in the Hyriinm (mostly absent in the Mutelince) ;
they are variable in number, generally only a few, and are located in the bottom
of the shallow beak-cavity, forming an irregular row parallel to the hinge-line, or
somewhat oblique. In some forms with deep beak-cavities, these scars are situated
on the inner side of the hinge-plate (behind the pseudocardinals and the inter-
dentum) .

The line of the mantle-impression is always simple, without a sinus poster-
iorly. The ligamental sinus lies on the margin of the shell behind the ligament,
over the posterior part of the lateral teeth (rarely more in front), and is always
small and shallow (longer than deep). (In the Mutelince, the ligamental sinus is
generally much deeper, with a sharp lower point.)

The characters of the soft parts of the Hyriince have been pointed out above.

The Genera of the Hyriinm.

The genera of the Hyriince have been hitherto differentiated according to
the shajie of the shell, the character of the beak-sculpture, and the character and
the sculpture of the hinge-teeth. According to Simpson (1914 pp. 1194 ff.) there
are seven of them in South America: Tetraplodori Spix {recte Castalia Lamarck);
Castalina Von Ihering; Castaliella Simpson; Callonaia Simpson; Hyria Lamarck;
Prisodon Schumacher; Diplodon Spix.

In addit ion, there are species lielonging to this group in Australia (and probably
also in New Zealand), which have been placed by Simpson in Diplodon, but have
lieen separated by him under the subgenus Hyridella Swainson. 1 have shown
(Ortmann, 1912) that the type of Hyridella {Unio australis Lamarck) actually is
closely allied 4o Diplodon, but that probably it is better to regard Hyridella as a
genus by itself.

ortmann: south American naiades.

461

The differences of the South American genera may be tabulated as follows:

a. Shell not alate, or only very slightlyalate behind, subelliptical, subovate, subrotund, or subtriangular.

b. Shell subelliptical, subovate, or subrotund, but not subtriangular. Beaks not much elevated,
with radial beak-sculpture, variously developed. Posterior ridge absent or poorly developed;
when present and distinct, the shell is elongated. Hinge teeth more or less smooth or dis-
sected, or corrugated, but without vertical parallel ridges. Interdentum very narrow.

Diplodon.

bb. Shell subtriangular (or subquadrate). Beaks more or less elevated and with rather deep beak-
cavities. With or without beak-sculpture. Posterior ridge well developed, defining a
distinct posterior slope. Hinge-teeth generally much dissected, and often with parallel
vertical ridges. Interdentum well developed, rather broad,

c. Sides of shell somewhat flattened, posterior ridge moderate, posterior slope elevated in the
middle and slightly alate (thus the shell becomes subalate). Hinge-teeth smooth, crenu'

lated, or with parallel ridges Caslalina.

cc. Sides of shell more or less convex, posterior ridge sharp, posterior slope truncated, not
elevated (or only so in the young). Hinge-teeth with parallel ridges.

d. Radial beak-sculpture present.

e. Beak-sculpture strongly developed, extending over a large part of the disk. Surface

not concentrically sulcated Castalia.

ee. Beak-sculpture short anteriorly, extending farther on the disk posteriorly. Surface

covered with strong, even, concentric ridges (sulcated) Castaliella.

dd. Beaks without distinct sculpture, and surface of shell smooth Callonaia.

aa. Shell subrhomboidal or subtriangular, strongly alate behind, slightly alate in front.

b. Beaks with strong radial sculpture. Posterior ridge moderately developed Hyria.

bb. Beaks without sculpture. Posterior ridge strongly developed, sharp and high Prisodon.

From the expressions used in distinguishing these genera, it appears that
Diplodon is a rather composite genus, containing shells of various, more or less
indifferent types, while the other genera represent more marked and peculiar
shapes, with Caslalina forming, to a degree, a connection between Diplodon and
the rest. This indicates that Diplodon includes the more primitive and generalized
forms of the subfamily, while the others are more specialized and advanced. In
a general way, this is supported by the investigation of the anatomy. However,
it is much to be regretted that I am unable to give any information as to the struc-
ture of the soft parts of the genera Castaliella and Callonaia. The former is. en-
tirely unknown to me, and of the latter I possess only two odd valves. Also of
Prisodon I only have the soft parts of a young individual. Better material is at
hand to represent the other genera.

Generally speaking, the anatomy of all these forms is rather uniform; yet
there are certain characters in which they differ, and some of these, indeed, indi-
cate a gradual advance in the structure.

462

MEMOIRS OF THE CARNEGIE MUSEUM.

One of these characters has been noticed long ago, and has been discussed in
detail liy Von Ihering. This is the tendency manifested in certain forms to close
the branchial opening in front by a connection of the mantle-margins, which is
a character entirely wanting in the Unionidce, but it is found occasionally in both
subfamilies of the Mutelidw {Hyriincc and Mutelinm). But numerous members
of this family do not have this character, and thus it is evident that the absence
of this mantle-connection indicates a primitive condition, while its presence is a
more advanced stage, expressing the tendency to transform the branchial opening
into a closed tube (siphon).

According to Von Ihering (1898), the mantle-connection closing the branchial
opening anteriorly is always iiresent in Castalia. 1 have been able to confirm
this only partially. Of typical s]iecies of Cadalia 1 possess the soft parts of six
specimens of C. aciUicosta: they show the liranchial opening closed with one ex-
ception, where it is open. In C. undosa, a somewhat aberrant type. Von Ihering
describes the branchial as normally closed, but he says that in four out of twenty-
one specimens examined the mantle-connection was missing.^ Of this species I
have five specimens with soft parts, of which three have the branchial closed, while
it is open in two of them. In one of the latter, however, a small one, it may be
torn. It must be admitted that also in other cases this connection of the mantle
may have been torn in life or in preservation.'’ And thus it might be that normally
this mantle-connection is present in Castalia.

Castalina, according to Von Ihering, varies in the development of this character
in the different species and individuals. He has investigated two males of C.
nehringi, and one had the branchial closed, the other open. In eight females ex-
amined liy myself, I found the branchial open. In C. martensi Von Ihering found
the branchial opening in most cases closed. I have no soft parts of this species.
But in two specimens of C. psammoica, which I possess, one has this opening dis-
tinctly closed in front, in the other this is not the case.

Nowhere else within this subfamily has this character been found. Castaliella
and Callormia may possess it, but their anatomy is unknown. Of Hyria and Prisodon

* In Von Iliering’s description (1893, p. 86) there is a very singular mistake. He speaks of the
mantle-connection “ Bruecke ” between the two “ siphons,” while he undoubtedly means the con-
nection in front of the branchial “ sii)hon.” This is the one which is sometimes missing. The absence
of the connection between the anal and l)ranchial openings would be something unheard of; in fact,
has never been observed bj^ me in any member of the family.

We must recall that similar variations or mutilations with regard to the mantle-connection which
separates anal and supra-anal oi)enings in the Unionidoi are well establishctl, for instance in the genus
Fuscunaia.

ortmann: south American naiades.

463

it is positively known that no such mantle-connection exists, and the same is the
case in all investigated species of Diplodon.

Thus it is seen that Castalina and Castalia (and possibly also Castaliella and
Callonaia) form a group within the subfamily, which is not only specialized in the
shape of the shell, but also in the tendency to close the branchial opening an-
teriorly, As Castalina forms a transition from Diplodon to Castalia in the shell,
so it is transitional also in this latter character, and thus it is perfectly clear that
Castalina and Castalia cannot be primitive types, and other peculiar characters,
which they possess, probably also should not be regarded as primitive.

This is especially true of the parallel ridges vertical to the edge of the lateral
hinge-teeth (also sometimes present on the pseudocardinals). This character is
best developed in Castalia (Plate XXXIX, figs. 8c, 8d), but Castaliella is also de-
scribed as showing traces of it, and Callonaia has it. But here again Castalina
is transitional toward Diplodon. Castalina nehringi has, according to Von Ihering,
hinge-teeth obliquely corrugated, but never with parallel ridges vertical to the
edge. This is quite correct, as far as concerns the laterals; but some of my
specimens show that the furrows of the cardinal teeth occasionally may be nearly
parallel, thus producing nearly parallel ridges. In Castalina niarte7isi vertical
ridges are present and well-developed, at least in the anterior section of the lateral
teeth, but young specimens do not show them. My specimens of Castalina psam-
moica have the vertical ridges poorly, or not at all, developed.

This sculpture of vertical ridges upon the hinge-teeth in Castalia and the
allied genera has been much discussed in the literature. It is well known that
Neumayr believed that these ridges are homologous to similar structures seen
in the Trigoniidae, and that Castalia should be directly connected with Trigonia,
and this idea has been taken uj) by other authors. However, Von Ihering (1893
p. 85) has pointed out that this is incorrect. From the above considerations we
now know, that there undoubtedly is a genetic series, which leads from Diplodon
through Castalina to Castalia, with the beginning at Diplodon, and the end at
, Castalia; and, consequently, the vertical ridges on the hinge-teeth in Castalia are
not an ancestral character, inherited from Trigonia, but are a new acquisition,
marking an independent, comparatively high development of the Naiad-hinge.

Hyria and Prisodon may, or may not, possess these vertical ridges on the hinge-
teeth, If present {Prisodon alatus, PI. XL, figs. 2c, 2d), they are seen chiefly in

This idea has also been suggested by Pompeckj (Gegen Steinmann’s Geologische Grundlagen der
Abstammiingslehre, iin 3ten Jahr.-Ber. Niederssechs. Geol. Ver., 1910, p. 12) in criticizing Steinmann’s
attempt at a polyphyletic derivation of the Naiades from various Trigoniida;. Odhner (1918, p. 577,
footnote) also considers the sculpture of the teeth in Castalia as a secondarily developed character.

464

MEMOIRS OF THE CARNEGIE MUSEUM.

old specimens, while they are lacking in the younger ones, but old specimens also
may not show them. Thus these two genera should be comjiared in this respect
with Castalina, and it is cpiite possible that they have here their roots, the shape
of the shell of Castalina also indicating in its elevated posterior slope the “alate”
shape of Hyria.

Thus we see that the genera Castalina, Castalia, and probably also Castaliella
and Callonaia, form a series descended from Diplodon, characterized by peculiar
tendencies in the shell and. in the soft parts. The shell tends to assume a more
or less triangular shape, with high beaks, with a strong posterior ridge, and a trun-
cated posterior slope. The hinge-teeth tend to develop vertical ridges, and the
interdentum becomes wider. The soft parts tend to close the branchial opening
in front. All the rest of the soft parts, however, remain here in a comparatively
primitive state, being identical with, or standing very close to, the most primitive
type of Diplodon in the interrujited character of the septa of the gills (See below).
It also should be mentioned that the glochidia of these forms, where known, as in
Castalina and Castalia, possess the same shape and the peculiar hooks observed in
many species of Diplodon, but not in all (See below).

Apparently a side-branch of this series is formed by Hyria and Prisodon.
Here the ^‘alate” character of the shell, slightly indicated in Castalina, is empha-
sized, and develojied to an extreme degree; the hinge-teeth have a slight tendency
to develop vertical ridges, exactly as in Castalina: but in the anatomy" these forms
do not show the tendency to close the branchial opening in front. Also in the
rest of the soft parts they remain upon the jirimitive Z)fp/odon-stage.

We may express the affinities as follows:

Callonaia Castaliella Prisodon

Castalia

Castalina

1

Diplodon

The genus Diplodon yet remains to be discussed. We have referred to it
above as the most jirimitive type within the subfamily, and there are undoubtedly
a number of species contained in it which have a primitive structure. But there
are others which differ from them. The first question, however, to be considered
is which characters we should regard as primitive.

As far as the shell is concerned, there is considerable variety within this genus.

ortmann: south American naiades.

465

Most of the species are subelliptical or subovate, and moderately elongated; but
others approach more or less the suborbicular shajic. For the latter, Simpson
has introduced the subgenus Cydornya, and it is true that these form a rather well-
defined group, although there are certain species which seem to be intermediate.

The subquadrate or subtrigonal shape of the Castalia-group is unknown in
Diplodon, and the well-defined jiosterior ridge characteristic of this group is very
rarely found; the species, which have it, are always rather elongated (not trigonal),
so there cannot be any mistake about them.

The hinge-teeth are also variable, but generally represent, or are easily com-
pared with, the normal type described above (p. 459). There are one or two
pseudocardinals and two laterals in the left valve, and two jiseudocardinals and
one lateral in the right valve. The pseudocardinals may be stum])y, or more or
less compressed. They, as well as the laterals, may be smooth, or have corruga-
tions, or may be split and dissected. They never have parallel ridges.

The beak-sculpture also varies a good deal. It is always radial, but the
ridges composing it may be shorter or longer, finer or heavier, smooth, or dissected
by growth-lines, granular, and sometimes they may be irregular.

But all of these characters of the shell are connected with each other by nu-
merous transitions, and it is impossible to say that any one type is more primitive
than another. It is also extremely difficult to arrange the species of Diplodon into
groups according to these features. Simpson (1914 pp. 1225, 1228) has divided the
genus into subgenera and subordinate groujis, and has attempted to condense the
results in the shape of a ‘‘key,” but this key is practically worthless, of which fact
anyone may convince himself, when he tries to use it for the identification of a
species.

However, when I studied the soft parts of the various species at my disposal,
I discovered that there are rather well-marked and ap]iarently important diaraders
shown by the anatomy, especially by the structure of the marsupial part of the inner
gill of the female, while the rest of the anatomy is the same in all species.

The interlaminar connections of the gills are extremely weak in the male of
Diplodon, and in the outer gill of the female (which has the same structure as in
the male). They correspond to the description given by me for Hyria and Castalia
(Tetraplodon) (Ortmann, 1911a, pp. 115, 117). (See the figures of gills of various
species on Plates XLV, XLVI, XLVII; also sections of the gills on Plates XLVII
and XLVIII). These interlaminar connections are few and scattered over the
face of the gill, and do not form distinct septa (thus resembling the condition seen
in the Margaritanidce). But in the marsupial part of the inner gill of the female.

466

MEMOIRS OF THE CARNEGIE MUSEUM.

these connections become more frequent, are heavier, and stand closer together.
Very often we see short connections grouped together either in a reticulate form,
or in rows parallel to the gill-filaments, fe., vertical to the edge of the gill (See text-
figs. 1 and 2 f, on pp. 455-6), and it should be noticed that precisely the same ar-
rangement is also seen in the Australian Hyridella (Ortmann, 1912, pp. 100-103,
fig. 1). The connections of adjoining vertical rows may alternate in an irregular
way, and may thus form irregular transverse or oblique rows. Towards the edge of
the gill, and near the base, the connections are often somewhat elongated, stand
closer together in the same row, and thus a more distinct arrangement into vertical
septa is brought about, separating more distinct water-tubes (ovisacs), which, how-
ever, are laterally connected with each other by the interruptions of the septa
(See PI. XLV, figs. 16, 26, 3; PI. XLVI, figs. 1, 2, 3, 4, 6, 7a, 76; PI. XLVII, fig. 1).

There is no doubt that the arrangement described above is primitive, for it
most closely aiiproaches the condition seen in the male gill. The marsupial struc-
ture is simply brought about by a more frequent and heavier development of the
interlaminar connections of the male, which approach each other, and partly ar-
range themselves in vertical rows.

A step in advance is observed when the vertical rows prevail over the reticulate
arrangement, and extend over the whole marsiijiial portion of the gill. Such cases
are found, and in them the interlaminar connections may be shorter or longer,
but they always stand close together in each row, so that we have all through the
marsupium the appearance of distinct septa, which, however, are perforated by
holes, so that the water-tubes (ovisacs) communicate with each other; this is most
evident in Diplodon piceus (PI. XLVI, fig. 2).

Furthermore in a few species (D. decipiens, PI. XLV, fig. 46, and D. mogymirim,
PI. fig. 5c; see also PI. XLVII, fig. 7, and PI. XLVIII, figs. 2a, 26), I have observed
a further advance in this structure. Here the septa become solid, the interruptions
are missing, and each septum runs from the base of the gill vertically towards the
edge, exactly in the manner which we know to be the characteristic condition in the
Unionidce. Well-defined water-tubes, which do not communicate, are thus formed.
The specie^ which show this structure, have in the marsupial part of the gill
hardly any indications of interruptions of the septa. In the non-marsupial por-
tions, as well as in the outer gill of the female, and both gills of the male, the usual
structure of Diplodon is present, showing few and scattered connections. This
reveals that the solid septa of these species of Diplodon represent the highest stage
in the marsupial development in this genus, and that this feature should not be
considered as homologous to the se]ita of the Unionidce. It is analogous, and has
been independently acquired.

ortmann: south American naiades.

467

There are other differences in the marsupium of the species of Diplodon, and
these depend upon its location within the gill. In the Australian Hyridella nearly
the whole inner gill is marsupial, only very small jiortions at the anterior and
posterior ends remaining non-marsupial, a very common condition in the Unionidce
where the whole outer gill may be marsupial. In Diplodon we sometimes observe
a similar arrangement (0. piceus, Plate XLVI, fig. 2), but generally a more con-
siderable part at the anterior as well as at the posterior end of the gill is non-
marsupial, so that the marsupium is restricted to the middle portion (about half)
of the gill. The marsupial part may become still more reduced in size, and may be
located not in the middle of the gill, but more toward the front, or toward the pos-
terior end. In either case the posterior or anterior half of the gill is nearly or
quite non-marsupial. It should lie noted that I have observed this shifting of
the marsupial part only in cases where the marsupial structure is comparatively
primitive, with the interlaminar connections arranged in a reticulate way or as
interrupted septa, but never in forms with solid septa, where the marsupial part
always extends over a large section in about the middle of the gill. (Compare
figures on PI. XLV, XLVI, XLVII.)

All these differences described in the structure and location of the marsupium
are constant within the species. In some forms, indeed, my material is rather scanty;
but in quite a number of other cases I have a sufficient number of individuals
with soft parts, and I have invariably found that all females of the same form and
from the same locality agree with each other, with the only qualification that in
young females the marsupium is generally less extended, and occupies a smaller
section of the gill (Conqiare PI. XLVI, figs. 56 and 5c; PI. XLVI, figs. 7a and 76).

It should be emphasized that these niarsupial differentiations are only found
within the genus Diplodon. The other genera of which I have anatomical material
stand generally upon the stages which I have described as the more primitive in
Diplodon. Firstly the structure of the marsuiiium is of the reticulate type in
the middle, with more or less development of interrupted sejita towards the edges
of the gill in Hyria (Ortmann, 1911a, p. 115) and Castalia undosa {ibid., p. 117),
while in Castalma nehringi the structure showing perforated sejita jirevails. Cas-
talia acuticosta seems to agree with C. undosa, but my material consists of only two
gravid, rather young specimens, in which the structure cannot be clearly seen on
account of the mass of glochidia filling the marsupium. Secondly the location of
the marsupium in Hyria and Castalia is in the middle of the gill (one-fourth at
anterior end, less than that at posterior end, non-marsu])ial). In Castalina nehringi
(Plate XLVII, fig. 2) the marsupium has moved a little more backward with about

468

MEMOIRS OF THE CARNEGIE MUSEUM.

the anterior half of the gill, and less than one-fourth of the posterior end non-
marsupial.

Thus it is clear, that, while the Castalia-Hyria-gYO\\\) of genera in the shape of
the shell and the hinge-teeth, and the Castalia-grow]) in the conformation of the
branchial openings represent a higher specialisation, in the rest of their anatomy,
and chiefly in their marsu])ial characters, they all remain very close to the primitive
Z)ip/od'on-type.

The Glochidia.

(See Text-figure 4.)

In conclusion I may say that I have found differences in the glochidia within
the genus Diplodon. That certain South American species of Ihiio” have glochidia,
was first announced by Lea in the case of U. peculiaris and U.firnius (Lea, Obs., XII,
1869, PI. 34, figs. 80, 82; first published in 1868). He describes and figures them as
subtriangular in outline, oblique, or upright, the ventral margin with a point, and in
the text he says that they are ^‘furnished with hooks.” However, he neither
describes nor figures these hooks.

Von Ihering (1893, p. 47) states that hooks are missing in all of the South
American species examined by him, but that the larvae are true glochidia.

This is aliout all we have known hitherto about the larval form of the South
American “Unios” = Hyriince. I haA^e now found that the normal shape (out-
line) of the glochidia of Diplodon, and of the subfamily Hyriince, is as described by
Lea, namely subtriangulai', with a point on the lower margin (see fig. 4). In addi-
tion, I have found that some species actually possess a ‘diook;” but this hook is
entirely different from the one so well known in the European genus Unio and in
the Anodontince of Eurasia and North America. The very fact that Lea mentions
the hook in a kind of perfunctory Avay, not calling attention to its iieculiar features,
suggests that he never saw the real hook, and that he simjily took the point of the
lower margin of the glochidium for it.

This Hyriine hook (Fig. 4, b, c, d, e, f, h, k, I, m) differs entirely from the Ano-
dontine hook. The latter is triangular, attached by a broad base to the point of
the lower margin, and carries upon its upper surface a number of fine spinules.
The Hyriine hook is long and narrow, spiniform, with very narrow base, articulated
to the point of the lower margin, and without any spinules on the upper face, and
furthermore has a ]ieculiar S-shaped curve.

It is jierfectly clear that this hook is different from that found in the genus
Unio and the Anodontince. Functionally it may serve the same purpose, that of

ortmann: south American naiades.

469

forming attachment to fishes, but we have not the slightest direct evidence of this,
and Von Ihering directly questions it (1. c., p, 47). Morphologically this organ has
been independently developed.- It is analogous to the Anodontine hook, but not
homologous.

Fig. 4. Glochidia of Hyriince, enlarged fifty times (drawn from photographs).

a. Diplodon hasemani Ortmann (Rio Giiapore), from specimen No. G. Cat. No. GI.5857.

b. D. imitator Ortmann (Santa Maria), from specimen No. 22. Cat. No. G1.9248.

c. D. simillimus Ortmann (Morretes), from specimen No. 2. Cat. No. G1.9250.

d. D. vicarius Ortmann (Aqua Qiiente), from specimen No. 9. Cat. No. G1.9251 (shell of this specimen

figured on Plate XXXVI, fig. 2).

e. D. decipiens Ortmann (Serrinha), from specimen No. 10. Cat. No. G1.9253.

/. D. paulista (Von Ihering) (Mogy Mirim), from specimen No. 10. Cat. No. G1.92.5G.

g. D. charrrianus (D’Orbigny) (Santa Isabel), from specimen No. 5. Cat. No. GI.08GI.

h. D. piceus (Lea) (Uruguayana), from specimen No. 1. Cat. No. G1..58G2.

i. D. hildw Ortmann (Cachoeira), from specimen No. c. Cat. No. G1.58G4.

k. D. mogymirim Ortmann (Mogy Mirim), from -specimen No. 48. Cat. No. 61.92G0.

l. Castalina nehringi Von Ihering (Salto das Cruzes), from specimen No. 1. Cat. No. G1..5119.
m. Castalia acuticosta Hupe (Rio Guapore), from specimen No. 1. Cat. No. 01.5112.

I have said that only some HyriinoB have this hook. I found it in the case of
several species of Diplodon, in Castalina nehringi, and Castalia acuticosta, and it

470

MEMOIRS OF THE CARNEGIE MUSEUM.

should be mentioned at this point that these hooks appear only in the fully developed
glochidium, while in younger, immature giochidia they are lacking.

But it seems that in certain species of Diplodon the hooks are always absent,
and in these cases the giochidia fully resemble the figures given by Lea. Of course,
in certain cases it is hard to decide whether the giochidia are fully developed, or
whether the absence of hooks may be due to the immature condition of the indi-
vidual. However, I have a case in which I believe I am justified in thinking that
the giochidia, when fully developed, have no hooks. In three gravid females of
D. charruanus, the giochidia were all alike, and no hooks were observed (Fig. 4g).
One of these apparently was discharging, and thus we should expect mature giochi-
dia (unless this were a case of premature discharge).

In a few other cases I am sure that mature giochidia have no hooks, since
they are surrounded around the whole lower edge, with a margin, which possibly
represents the first beginning of the permanent shell of the adult (Fig. 4a, i). In
other grou])s of Naiades, with one exceiition {Anodonta imhecillis Say of North
America), nothing of the kind is known in giochidia, as long as they remain within
the marsipiium, and also in these South American forms, when immature, the
giochidia do not jiossess this margin. I never have seen a trace of hooks here.
The margin has much the apjiearance of that formed in young North American
Unionidw, after the parasitic stage on fish.

That in these cases the shell should appear at so early a stage, when the larva
is still within the marsupium of the mother, is indeed remarkable, and possibly
points to the conclusion that the modes and conditions of embryonic develop-
ment in the Hyriince differ considerably from those of the Unionidce.

If the aliove observations are correct, we would have three types of giochidia
in the genus Diplodon: (1) Of triangular shape, with hooks; (2) Of the same shape,
without hooks; (3) Of the same shape, and without hooks, but with a margin around
the lower edge, which obliterates the triangular shape.

Whether the second gi-oup is real, or only due to incomplete observation, or
passes finally into the third, remains to be seen. At any rate, it is very desirable
that close attention should be directed to this cpiestion in future work on South
American Naiades.

The size of the giochidia is comparatively large, varying from 0.20 to 0.35 mm.,
which might be called a good medium size in comparison with the giochidia of the
Unionidce.

I regret that my observations on the giochidia of the Hyriince are not more
satisfactory. There are indications of important differences, but for the present

ortmann: south American naiades.

471

we must be satisfied with having pointed out this fact, and with hinting that per-
haps further knowledge of the glochidia may furnish at least some criteria for a
classification of the species of Diplodon. The same remark holds good of the mar-
supial structure. In reference to this we possess a little more knowledge, but it
is still too scanty to make an attempt to use it for classification. The genera with
the specialized shells of the Castalia- and Hyria-gmiips are well defined, but their
anatomy and their glochidia do not present any remarkable differentiations, except
the structure of the branchial opening in the Castofia-group. On the other hand,
it appears that the genus Diplodon may finally prove to be composed of an aggrega-
tion of more varied forms than heretofore supposed, and that the distinctive char-
acters of these are found chiefly in the marsupium and the glochidium.

We may perhaps distinguish within the old genus Diplodon a type, which we
might regard as having the more primitive features, such as a marsupium com-
posed of interrupted or reticulated septa, occupying the whole or nearly the whole
of the inner gill, and probably one of the types of glochidia above described. But
it is hard to say, which form this is. I am inclined to regard the hooked glochidium
as the more primitive form. The margined glochidium certainly represents a
more advanced type. It might finally be possible to split up this genus into smaller
genera, one of which should contain these primitive forms, the others to include
forms more advanced in regard to their marsupial structure and glochidia. But
in the present state of our knowledge this step cannot be taken, and we must be
satisfied with an arrangement of the species based upon the characters of the
shells.

The Species of the Hyriin^.

Genus Diplodon Spix (1827).

Diplodon Spix, 1827, p. 33 and plate 26. — Diplodon Simpson, 1900, p. 872; 1914,

p. 1224.11

Type of genus: — Diplodon ellipticuni {ellypticum err. typogr.) Spix, 1827, PL
26, figs. 1, 2. (Same type given by Simpson, 1900.)

The Subdivisions of the genus Diplodon.

Simpson (1914, p. 1225) has made an attempt to divide this genus into sub-
genera and groups. But he also included in it Australian species under the sub-
genus Hyridella Swainson, which we now regard as a separate genus, and an African

Diplodon, as accepted by H. & A. Adams (1858, p. 497) as a subgenus of Unio, is an entirely
heterogenous association of species, including forms from all over the world.

472

MEMOIRS OF THE CARNEGIE MUSEUM.

form (Subgenus Lcevirostris Simpson), which we may with considerable confidence
assume does not belong here, until the contrary has been positively demonstrated.

Thus restricted, the genus Diplodon from our point of view falls in Simpson’s
arrangement into two subgenera, Diplodon (Simpson, 1914, p. 1226) and Cyclomya
(Simpson, 1914, p. 1278), each subdivided into groups. The characters of the
shells of these groups run into each other very insensibly, and are found in various
combinations. The subgenus Cyclomya is somewhat better defined, inasmuch as
the rounded outlines of the species assigned to it contrast rather markedly with
most of the elliptical, somewhat elongated shells, assigned by Simpson to typical
Diplodon. But there are intergrades even here. Typical Cyclomya (of the fune-
hraUs-type) is irregularly circular, with rounded angles (obscurely pentagonal),
and has a short and high shell, with the greatest height situated in the middle of the
shell, at about the middle of the ligament. Simpson included in it as species
fontainianus, and gratus, which 1 regard only as higher and shorter forms belonging
to Diplodon. They have the greatest height of the shell not in the middle, but
more posteriorly, behind the ligament, and thus their outline is distinctly oblique
and subtrapezoidal, although approaching the rounded shape.

Simpson, indeed, says in addition, that the beak-sculpture in typical Diplodon
consists of ‘Tmbroken ridges,” and that in Cyclomya it is “irregularly radial.”
In both cases this holds good only for certain species, and cannot be used as a
generally distinguishing character.

Subgenus Diplodon Simpson.

Simpson, 1900, p. 873; 1914, p. 1226.

Shell more or less elongated; elliptical, ovate, or subtrapezoidal in outline;

I*

when short, more or less angular and distinctly oblique, with the greatest height
behind the ligament, but not subcircular with the greatest height under the ligament.

The arrangement into groups, as here given, does not rest upon that of Simpson.
It is largely made to suit mj^ material, and does not claim to be final. We may ex-
jiress the essential differences in the following key, but with the distinct under-
standing that there are transitions between the groups, which are hard to place.

Key to Groups in Genus Diplodon.

a. SJiell straight, not oblique, i.e., the longest axis is nearly parallel to the ligament.

b. Beak-sculpture covering a considerable part of the disk; ridges, chiefly the posterior, rather

heavy Group of D.' hijlccus.

bb. Beak-sculpture more or less developed, rarely covering a considerable part of the disk; ridges
not very heavy and rather uniform.

ortmann: south American naiades.

473

c. Beak-sculpture well-developed, fine; ridges cut up into fine nodules.. .Group of J). granosus.
cc. Beak-sculpture restricted to the region of the beaks; ridges fine, simple, not granular.

d. Shell rather compressed and flattened upon the sides, subtrapezoidal in outline, and

not distinctly pointed behind Group of I), chilensis.

dd. Shell slightly or not at all compressed, convex upon the sides, mostly subelliptical or
subovate in outline, more or less pointed behind.

e. Shell rather elongate, not high, subelliptical or subovate. . .Group of I), charruanus .

ee. Shell shorter and higher, subovate or sidjtrapczoidal Group of D. lacteolus.

aa. Shell distinctly oblique, with the longest axis forming an angle with the line of the ligament. Outline
subovate or subtrapezoidal, sometimes rather high and short Group of D. ellipticus.

1. Group of Diplodon hylceus.

Shell subelliptical, subovate, or subtrapezoidal, more or less pointed posteriorly,
straight, not distinctly higher behind, nor oblique. Beak-sculi)ture well-developed,
covering a considerable (but variable) part of the shell; bars rather heavy; those
upon, and immediately in front of, the indistinct posterior ridge of the shell are
heavier and longer than the rest; one or two of the median bars joining at their lower
ends; sometimes the bars are somewhat nodulous (but not granular).

The beak-sculpture is the most essential feature of this group. Additional,
but subordinate, characters are found in the hinge-teeth, chiefly the larger ones in
each valve, which are triangular, rather thick, ragged, but not compressed. Often
there are two pseudocardinals in the left valve.

I have seen the female soft parts of two species, D. hasemani and trijidus.
In both the marsupial part of the inner gill is located in the middle section and
has interrupted septa. The former species has the margined type of glochidiuni.
It seems that this group is not very primitive.

1. Diplodon hyl^us (D’Orbigny).

Unto hylcea D’Orbigny, 1835, p. 36; 1843, p. 607, PI. 69, figs. 8, 9.

Unio hylceus Sowerby, XVI, 1868, PI. 93, figs. 506 a and h (poor hgures).
Margaron {Unio) hylceus Lea {pro parte), Syn. 1870, p. 31.

Unio hylceus Von Martens, 1894, p. 164.

Diplodon hylceus Simpson {pro parte), 1900, p. 884; 1914, p. 1274.

Type-locality. — “Rio Palometas, Rio Pari, and Rio Tucabaca, in the provinces
of Santa Cruz de la Sierra and Chiquitos, in Bolivia.”

I have been unable to locate the first two rivers; a Rio Tucaraca, in the Chi-
quitos country (Prov. Santa Cruz de la Sierra in Bolivia), runs to the Paraguay
River, and I have no doubt that this river was intended, and it is advisable to take
this as the type-locality.

474

MEMOIRS OF THE CARNEGIE MUSEUM.

Other lucalities. — Von Ihering (1893, p. 119) gives this species from Rio Para-
guay, but also upon D’Orbigny’s authority,^" from the Amazon-drainage. Von
Alartens {1. c.) reports it from Paraguay. No other localities have been hitherto
cited.

New locality. — Rio Paraguay, Sao Luiz de Caceres, Matto Grosso, Brazil,
(J. D. Hasenian coll.. May 25, 1909). One specimen.

Distribution. — The new locality is not very far from the type-locality (Rio
Tacaraca), about 400 kilometers up the river, and demonstrates that this species
belongs to the upper Paraguay drainage in Bolivia and western Brazil. According
to Von Martens it goes down this river to Paraguay.

D. hylwus and guaranianus have been united by Lea, Sowerby, and Simpson,
but 1 think that this is not correct, and that I have both species before me, agree-
ing well with D’Orbigny’s remarks about them. The real D. hylwus is a larger
shell, with the lower posterior end somewhat more produced, and with differences
in the beak-sculpture, which extends over a larger section of the shell, and con-
sists of rounded bars, which are somewhat irregular, and are rugose or slightly
nodulous. The figure of D’Orbigny (Fig. 8) shows the sculpture very well.

I have only a single individual of this species, without soft parts, and have
drawn the following description from this.

Description of Shell. — Shell comparatively small, moderately solid, subovate
or subtrapezoidal, slightly higher behind. Height 04 pr. ct. of length. Dorsal
margin very gently curved, passing into the posterior margin in a blunt, rounded
angle. Posterior margin obliquely descending, emarginated, but this emargina-
tion undoubtedly is an individual feature, since the growth-lines indicate that it was
not present, when the shell was younger. Lower jiosterior angle slightly pro-
duced, but rounded. Lower margin with its lowermost part jilaced at between
two-thirds and three-fourths of the length of the shell (from anterior end), curving
up behind. In the anterior iiortion it slopes upward in an almost straight line,
finally curving up into the anterior margin; thus the shell appears slightly narrower
in front than behind.

Valves rather flat (ii; this respect my specimen differs from the original hylcEUS,
which is more convex), gently and rather uniformly convex, with the umbonal
(l)osterior) ridge weakly marked, and indicated chiefly by a shallow radial de-
pression 111)011 the posterior slojie, which forms the emargination at the posterior
margin, and makes the posterior ridge appear slightly biangulate towards the

• He says that all forms reported by D’Orbigiiy from Bolivia are from the Amazon-draiiiage. In

this particular case this is not correct, since the Rio Tucaraca of Bolivia drains into the Paraguay.

ortmann: south American naiades.

475

posterior end. Diameter of shell 33 pr. ct. of length. Beaks not swollen, and
not prominent, located at about one-fourth of the length from the anterior end.

Beak-sculpture strongly developed, covering about half of the disk, the longest
bars (near the umbonal ridge) are about 20 mm. or more long. There are about
sixteen or seventeen radial bars, of which the ninth and tenth, and the eighth and
eleventh, unite in sharp angles, and between the ninth and tenth, there is a short
odd bar, which is indistinct on account of the erosion of the beaks. Posteriorly the
bars increase in length as well as thickness: the anterior bars are comparatively
sharp, narrow, but much injured in 1113^ specimen; the posterior bars are broader
and rounded. Upon the umbonal ridge, the radial bars are again finer, and upon
the posterior slope there are four or five additional fine bars, which are shorter,
and restricted to the upper part of the slope. On the lower part of the latter,
there are a number of fine, irregular, oblique wrinkles. The lower ends of the
radial bars are cut up, chief!}' near the umbonal slope, into irregular, low tubercles,
and traces of such tubercles may be seen near the lower margin of the shell.
No distinct lunula is seen in our shell, but this part of the shell is badly eroded.

Epidermis with numerous, irregular, concentric wrinkles, and traces of radial
lines. Color brown, much like that of D’Orbigny’s figure.

Hinge-line gently curved. Ligamental sinus over the posterior fourth (or a
little more) of the lateral teeth, which are gently curved, one in the right, two
in the left valve. Pseudocardinals directed obliquely forward and downward,
two in right valve, the anterior one narrow, low, and compressed, the posterior
one triangular, cut longitudinally into two parts. In the groove behind this
tooth, the hinge-line has two small denticles. In the left valve, there are two
pseudocardinals, the anterior subtriangular, slightly compressed and simple, fitting
into the groove between the two teeth of the right valve, the posterior one broader,
and cut into three parts. Leaving out of sight the comparatively stumpy and
double character of these teeth, their finer structure can only be regarded as an
individual characteristic.

Cavity of shell and beaks shallow. Nacre whitish (discolored in the cavity).
Anterior adductor-scar distinct, subelliptical; anterior retractor-scar separated
from it, small, round and deep; anterior protractor-scar connected with the ad-
ductor-scar. Posterior adductor-scar faint, subovate, with an upper triangular
process formed by the posterior retractor-scar. Alantle-line faint. Dorsal
muscle-scars in the cavity of the beaks.

476

MEMOIRS OF THE CARNEGIE MUSEUM.

Measurements.

Leugtli.

Height.

Diameter

Beaks.

Mv specimen

42 mm.

27 mm. =04 pr. ct. of L.

14 mm. =33 pr. ct. of L.

at 10 mm. =24 pr. ct. of L.

D’Orbifinv’s figure

43 “

25 “■ =58

10 “ =37

8 “ =19

Du., in text

45 “

00

40

Remarks: The chief characters of this species, so far as I am able to make
them out from the figure and description, and the single specimen at hand, by
which it differs from the other species of this group, are found in the general shape
(moderately elongated, slightly wider behind, with rounded posterior angle),
the lieak-sculpture, which covers a rather large portion of the disk, and the rough
character of the posterior slojie. The greater compression of our shell probably
is individua], and also the emargination of the posterior margin.

2. Diplodon guaranianus (D’Orbigny).

Uriio guarauiana D’Orbigny, 1835, p. 37; 1843, p. 608, pi. 69, figs. 10-12.

Type-locality. — Rio Parana, Itaty, Province of Corrientes, Argentina. No
other locality jireviously known.

New Localities. — Paraguay River, Corumba, Matto Grosso, Brazil (H. H.
Smith coll.). One specimen, juv. In swamps of Lambare, near R. Paraguay, Asun-
cion, Paraguay (J. D. Plaseman coll. March 31. 1909). One complete specimen,

6 left valves. Rio Paraguay, Sao Luis de Caceres, Matto Grosso, Brazil (J. D.
Ilaseman coll. May 25, 1909). One male, with soft parts, and seven specimens,
shells only.

Distribution. — Middle Parana above the junction with the Paraguay River in
Argentina and Paraguay, and Paraguay River through Paraguay as far as Matto
Grosso, Brazil.

I disagree with previous authors, and regard this species as being distinct from
D. liyUvus. My specimens answer very well to the description and figures of
D’Orbigny, and they differ from Ivylccus in their somewhat smaller size, in the more
distinctly truncated (more steeply descending) posterior margin,, forming a more
distinct lower iiosterior angle, but chiefly in the beak-sculpture, which consists
of a smaller number of bars, wliicli are heavier, chiefly posteriorly, and are not so
rugose. My siiecimens are also more swollen than the single individual of hylceus
at hand, but the latter is, as has been stated, probably exceptional in this.

The beak-sciiljiture is somewhat variable, chiefly in the length of the bars,
which are from 15 to 20 mm. long, and generally cover more than half of the disk,
but in the larger shells, sometimes less. There are twelve to fourteen radial bars,

ortmann: south American naiades.

477

of which the seventh and eighth, or ninth and tenth, unite in the middle of the
valve in a sharp angle; sometimes a short odd bar is found between these pairs.
The bars are rather fine and sharp in front, but those behind, near the umbonal
ridge become broad and rounded, their lower ends being irregular and indistinctly
tubercular (much less so than in D. hylceus) occasioned by concentric lines cutting
across them. These posterior bars also are distinctly longer than the anterior
ones. Sometimes there are a few additional fine bars near the beaks on the posterior
slope and below them a few oblique wrinkles.

The posterior end of the shell is also somewhat variable, but it never is rounded
and slightly biangular, as it is in D. hylcvus. The posterior margin is obliquely
descending, and the lower posterior end is bluntly pointed, the angle being more
or less prominent. The lower margin of the shell is evenly convex in some speci-
mens, in others it is more strongly convex a little back of the middle, forming a
blunt angle.

The specimen from Corumlia is young, and the beak-sculpture covers all of
the shell. It is also a little more compressed than the others, but agrees with them
in all other characters.

JMeasurements.

Length.

Height.

Diameter.

Beaks.

Asuncion, No. 1

21 nini.

14 nun.

= 07 pr. ct. of L.

10

nun. =48 pr. ct. of L.

at 4 nun. =19 pr. ct. of L.

Do. 2

27

18 “

= 07

14

“ =52

0.5 “ =24

Do. 3

28

19 “

=08

13

“ =40

7 “ =25

33.5 “

9‘;; “

-00

13

“ -39

8 “ -24

S. Luis d. C. (cf)

39

24 “

= 02

15.5

“ =40

8 “ =21

D’Orbigny’s fig

32

19 “

= 59

12

“ =38

8 “ =25

Thus my specimens are on the average slightly higher and shorter, and some-
what more swollen than the original, unless there are inaccuracies in the figure,
which is not impossible, since the figure is 32 mm. long, while the text gives the
length as 21 mm.

Anatomy. — The only specimen (S. Luis de Caceres) of which soft parts are
at hand, is a male, and has the structure typical of the genus. The anal open-
ing is slit-like, and about three-fourths of the length of the branchial. The
branchial opening has fine papilla;. The palpi are triangular and rather small,
the posterior margins are not connected. Inner lamina of inner gills connected
with abdominal sac. Gill-structure typical.

478

MEMOIRS OF THE CARNEGIE MUSEUM.

3. Diplodon hasemani Ortmanii, sp. nov.

PL XXXIV, figs. 1 to 4, shells; PI. XLVII, fig. 5, section of gills; text-fig. 4a (p.

469), glochidium.

Locality. — Rio Guapore, near Rio Sao Simao, Matto Grosso, Brazil (John D.
Haseman coll., July 20, 1909). Nineteen specimens investigated, all with soft
parts; sex of the three smallest not ascertained; the rest were males and gravid
females.

Type-set: Cam. Miis. No. 61.5857, twelve specimens, among them four males
and five gravid females, one with eggs, the others with glochidia.

Description of Shell. — Shell small (max. length 28 mm.), solid, swollen, short,
subovate, somewhat pointed behind. Diameter 67 to 72 pr. ct. of length, as against
59 to 68 pr. ct. in D. yuaranianus. Valves not gaping. Dorsal margin gently
convex, passing gradually or with a blunt angle into the anterior margin, which
is sometimes almost truncated. The posterior upper margin forms an obtuse,
rounded angle with the posterior margin, the latter descending obliquely, gently
convex, forming with the lower margin a rather distinct, but blunt posterior ter-
mination of the shell, which is little elevated above the base-line. Lower margin
gently and uniformly convex, passing in a regular curve into the anterior margin.
The anterior portion of the shell does not appear appreciably narrower than the
posterior, or very little so.

Valves convex, more so in older specimens, with a rather distinct, but rounded
umbonal ridge. Posterior slope subtruncated, a little compressed and elevated
in the middle. Greatest diameter of shell 42 to 55 pr. ct. of length (32 to 52 pr. ct.
in D. guaraniaiius). This greatest diameter located more forward toward the
beaks than in D. guaraniarms. Beaks somewhat swollen, but little elevated above
the hinge-line, located at 25 to 28 pr. ct. of the length. Beak-sculpture of the
hylceus-typc, strongly develojied, the posterior bars thicker and longer than the
anterior. They cover 10 to 15 mm. of the shell, that is to say, hardly half of it
in larger specimens. There are from fourteen to sixteen of them, and the ninth
and tenth generally meet at an acute angle, with sometimes a short odd one between
this pair. The anterior bars are sharp, the posterior ones broader and rounded,
but not so much as in D. guaranianiis. There are often three or four additional
finer bars upon the jiosterior slope, and some oblique wrinkles behind and below
them. The lower ends of the bars are but faintly cut up into tubercles. A lanceo-
late, rather short, lunula may be present in the larger specimens.

Epidermis with numerous, concentric, irregular stria3, finely lamellar in young

ortmann: south American naiades.

479

specimens, and in older specimens on the posterior slope and toward the lower
margin. Fine, obscure, radial lines all over the shell. Color rather uniformly
dark brown, but in young specimens there is a distinct indication of a yellowish
concentric band, or the region near the beaks may be of this color (suggesting the
color-pattern of D. trijidus).

Hinge-line gently curved. Ligamental sinus over the middle of the lateral
tooth, or a little farther back. Lateral teeth curved, one in right, two in left
valve. In one specimen in the right valve there is a distinct, but low, accessory
lateral below the normal one. As a rule two pseudocardinals in each valve. They
are not much compressed and not much elongated, but rather stumpy, and are
very much cut up, especially the posterior ones in each valve. The anterior
pseudocardinal in the right valve is more distinctly compressed, but may be very
small or even obsolete, and there may be a trace of a third pseudocardinal liehind.
The posterior pseudocardinal in the left valve is, very variable in shape and size.

Cavity of shell and of beaks moderate. Nacre in all specimens whitish.
Anterior adductor-scar deep, even in young specimens, irregularly rounded. An-
terior retractor-scar separated from it, small and deej). Anterior protractor-
scar united with the adductor-scar. Posterior adductor-scar less distinct, sub-
triangular, with a short upper process formed by the posterior retractor-scar.
Pallial line distinct. Dorsal scars few, located in beak-cavit}" and close to hinge-
plate.

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Figures.

1.. .

?

12.5

mm.

9

mni. =72 pr. ct. of L.

5.5

nim. =44 pr. ct. of L.

iit 3.5

mm. =28 pr. ct. of L.

2...

?

15

10

“ =67

7

“ =47

4

" =27

2a.,

cf

18

13

“ =72

7.5

“ =42

5

“ =28

4.. .

9

23

16

“ =70

11

“ =48

6.5

“ =28

PI. XXXIV, 6g. 3.

6.. .

9

25.5

17

“ =67

12

“ =47

7

“ =27

7.. .

cf

26

18

“ =69

14

“ =53

7

“ =27

9.. .

9

28

19.5

“ =70

14

“ =50

7

“ =25

PI. XXXIV, fig. 2.

10.. .

&

27.5

20

“ =72

15

“ =55

7

“ =25

PI. XXXIV, fig. 1.

I cannot discover any sexual differences in the shell.

Reynarhs. — There is no question that this species is very closely allied to
D. guaranianus, and it may be descrilied as a small, short, rather swollen D. guarani-
anus, with the posterior end of the shell a little sharjier, the posterior ridge a little
more distinct, and beak-sculpture less developed. Since I possess a good number
of specimens of D. hasemani, and also a fair number of D. guaranianus, I do not
doubt the specific distinctness, since the geographical distribution also differs.
D. guaranianus belongs to the Parana and Paraguay Rivers, while D. hasemani is
from the Guapore, tributary to the Amazons-system. However, a former con-
nection of these systems is indicated by the close affinity of the two species.

480

MEMOIRS OF THE CARNEGIE MUSEUM.

Anatomy. — I have before me many males and gravid females with soft parts.
It should be noted that the smallest gravid females are only 16 mm. long and that
this fact indicates that this is a small species, not growing to a larger size, as my
material shows, reaching the maximum length of 28 mm.

Anal ojiening short, slightly shorter than the branchial opening, the latter
very short, with few but distinct pa})illae. Palpi sulitriangular, lower margins
convex, posterior margins scarcely connected. Gills normal. Inner lamina of inner
gill entirely connected with abdominal sac. In the larger gravid females the
middle of the inner gill is marsupial, leaving a little less than one-fourth of the
gill lioth at the anterior and ]iosterior ends non-marsupial. In the smaller females
the marsupial part is smaller. The swelling of the charged marsipiium is moderate
and the gills are charged to near the edge. In the marsuiiial part the interlaminar
connections are arranged in interrupted vertical septa. Since no sterile females
are at hand, the exact arrangement could not be seen in a face view, but from sec-
tions (Plate XLVII, fig. 5) it is evident that it is very likely more or less reticulate.

Glochidia (Text-fig. 4n, p. 469) subtriangular and margined, without hooks.
Measurements, without margin: L. 0.30 to 0.31 mm.; H. 0.24 to 0.25 mm.; with
margin: L. 0.35 to 0.36 mm., H. 0.29 to 0.30 mm. The presence of eggs and
glochidia on July 20 should be recorded with respect to the breeding season.

Color of Soft Parts in Alcohol. — Foot blackish in its distal part, this color
separated in a sharp line from the light basal part. Rest of soft parts whitish.

4. Diplodon trifidus (Lea) (1860).

Diagram of Soft Parts {See Text-fig. 1, p. 455).

Unto trifidus Lea, Obs. X, 1863, PI. 44, fig. 295.

Diplodon trifidus Simpson, 1900, p. 884; 1914, p. 1272.

Type-locality. — ‘'Buenos Ayres.” Never reported again since its original
description.

New LoccUities. — Centre of Rio Guapore, near Rio Sao Simao, Matto Grosso,
Brazil (J. D. Haseman coll., July 20. 1909). Six specimens, all with soft parts.

In the three smallest the sex is uncertain (they have the male' structure) ; the others
are two males and one (the largest) a gravid female with eggs. Rio Guapore, Sao
Antonio de Guapore, Matto Grosso, Brazil (J. D. Haseman coll., July 31, 1909).
One male with soft parts.

Distribution. — I have doubts as to the correctness of the locality originally
given. Von Ihering (1893, p. 118) lists this species with those from the La Plata
drainage, but he rests entirely upon Lea. The latter received his single specimen

ortmann: south American naiades.

481

from D’Orbigny with the above locality; liat it is quite possible that a mistake
was made, since D’Orbigny also collected in the Amazon system in Bolivia, in
the general region, whence our specimens come. Nobody else ever re-discovered
this species in the La Plata, although frequent collections have been made. Our
material is beyond any doubt this species, and the locality is authentic.

The set, although containing only six specimens, comprises young and old,
males and females, and thus it is worth while to give a full descrijition.

Description of the Shell. — Of medium size, growing larger than any of the
preceding species (maximum length 58 mm.; the type is 42 mm., according to
Simpson), rather solid. Outline subelongated, subelliptical, or long-ovate, rounded
in front, pointed behind. Height 41 to 54 pr. ct. of length. Valves not gaping.
Dorsal margin straight in young specimens, very gently curved in older ones.
Anteriorly the dorsal margin forms an indistinct angle with the anterior margin
in young individuals; in old ones it passes into it gradually. Posteriorly the dorsal
margin forms a very obtuse angle with the posterior margin, or, in the largest
specimen, passes gradually into it in a gentle curve. Posterior margin obliipiely
descending, gently curved, meeting the posterior jiortion of the lower margin in a
blunt, but distinct, point, which is elevated above the base-line, but nearer to the
latter than to the line of the upper margin. Ventral margin gently and rather
regularly convex, its lowest point slightly behind the middle of the shell, ascending
in front and behind it. In front it curves up into the anterior margin. Thus the
shell has a long-ovate, almost subelliptical outline, with the anterior end only
slightly narrowed and rounded, and the ])osterior more tapering and bluntly pointed.

Valves moderately convex, convexity rather uniform all over the disk, but
strongest near the beaks and upon the umlional ridge, which forms a rounded, but
rather distinct, angulation running toward the posterior point. Posterior slope
somewhat compressed. Diameter 23 to 3G pr. ct. of length. Beaks not swollen,
hardly elevated above the hinge-line, located at 18 to 25 pr. ct. of the length. Since
the shell is more swollen towards the beaks, the latter appear depressed, chiefly
in old specimens. Beak-sculpture distinct and well develojied, consisting of about
sixteen radial bars; the anterior bars are shorter (about 5 mm. long), the posterior
much longer, chiefly upon the umbonal ridge, where they are 15 mm. long, or even
more. There are two systems of these bars; eight or nine anterior, and seven or
eight posterior bars. In the middle of the shell one or two pairs unite at a sharp
angle, and sometimes a short, odd bar stands between the innermost pair. The
anterior bars are sharp; the posterior ones are also rather sharp near the beaks,
but towards their lower ends they gradually become thicker and rounded, broaden-

482

MEMOIRS OF THE CARNEGIE MUSEUM.

ing and flattening, and finally disappearing. These broad bars are conspicuous
chiefly immediately in front of the umbonal ridge, and are distinctly seen even in
our largest specimens, where the beaks are greatly eroded. All these bars are
smooth, except for the concentric strise of the epidermis which cut across them.
In some of my young specimens there are a few oblique wrinkles upon the posterior
slope, but there are hardly any radial bars, except one, fine, and short, close behind
the broad bars of the umbonal ridge. There is a narrow and short lunula in front
of the beaks.

Epidermis smooth, but with numerous, irregular, concentric lines, which be-
come sublamellar upon the posterior slope and near the lower margin. There are
also numerous fine, irregular, but straight, radiating lines on the disk below the
beak-sculpture. The posterior slope has no radial ridges or furrows. Color of
epidermis dark to light green, with concentric bands of yellow-brown. The three
young specimens are light golden-brown towards the beaks, and light green towards
the margins; in the old specimens the green color prevails, and becomes quite dark,
but is interrupted by one or two lighter bands of brownish. No traces of color-
rays present.

Hinge-line almost straight or very gently curved. Ligamental sinus over the
posterior third of the lateral teeth. One lateral in the right, two in the left valve,
rather long, distinct, and in old shells gently curving downward behind. Pseudo-
cardinals very variable. Lea describes them as trifid in both valves, but this is
not always the case. Of our largest specimen it may be said that the right valve
has three teeth: the middle one the largest, directed obliquely downward and for-
ward, triangular, narrow above, broader below, and deeply longitudinally cleft
into three ridges; in front of it is an anterior, narrow, compressed tooth, which
is connected with the middle one above; behind the largest pseudocardinal is a
deep groove, followed by small, ragged elevation of the hinge-plate representing
the third tooth. The left valve has one large posterior tooth, which is ragged and
fits into the groove behind the middle tooth of the other valve (including the small
elevation behind it), and in front of it is a narrow lamellar tooth, fitting into the
space between the first and second teeth of the right valve. In the groove be-
tween these two teeth of the left valve are two low ridges, corresponding to the
clefts of the large tooth of the right valve. In our second largest specimen (No. 1)
the same general arrangement is seen, but the anterior tooth of the right valve is
very low, while the third (posterior) is more distinct and triangularly elevated.
The left valve has two teeth. All these teeth are much less ragged, and the clefts
of the middle tooth of the right valve are lacking, and also the corresponding ac-

ortmann: south American naiades.

483

cessory ridges of the left valve. Similar, but rather variable, conditions are seen
in the younger individuals and the specimen from S. Antonio. The rule is that
there are three teeth in the right, but only two in the left valve.

Cavity of the shell moderate, that of the beaks shallow. Nacre shining, snow-
white in my specimens, in two (No. 1 and the one from S. Antonio) with a faint
salmon blush in the cavity of the shell. Anterior adductor-scar deeply impressed
even in young specimens, irregularly rounded or broadly subelliptical. Anterior
retractor-scar located above it, separated from it, deep, small. Anterior protractor-
scar connected with it. Posterior adductor-scar distinct, but much less impressed,
subovate or subtriangular, with a rounded or triangular appendix above, formed
by the posterior retractor-scar. Pallial impression distinct. Dorsal scars five or
six, in cavity of beaks, i)laced irregularly, or in an oblique line.

There are no sexual differences in the shell.

Measurements (specimens from Sao Simao).

No.

Sex.

Length.

Height.

Diameter.

Beaks.

a

?

21.5 mni.

9 inin. =41 pr. et. of L.

5 mm

= 23 i)r. ct. of L.

at 4.5

mm. =21 pr. ct. of L.

h

d'

.’U

16 “ =47

10 “

= 29

7

“ =21

1

d'

50

30 “ =54

17 “

= 30

14

" =25

2

9

58

29 “ =50

21 “

= 30

14

“ =24

Lea’s figure

42

22 “ =52

15 “

= 30

7.5

“ =18

In Lea’s text there is a])parently an error with resiiect to the height, since the
measurement would yield the absurd figure of 89 ]w. ct. of the length. According
to the figure Lea had a specimen with the beaks a little more anterior than any of
mine, but otherwise rather closely agreeing with them.

Remarks. — This species is related to those of the hykeus-group, chiefly to
D. guaraniarius, but it is easily distinguished by its long-ovate shape, with pointed
posterior end, and by the beak-sculpture, which, although of the same type, does
not extend so far upon the disk. A good character is furnished also by the color
of the epidermis and its concentric bands, though we have seen that this ]iattern
is at least indicated in young D. hasemani. Lea compares D. trifidus with D.
burroughianus and parallelopipedon, but there hardly is any close relationshiji to
the former, and but a superficial resemblance to the latter. I think it belongs to
the hylwus-group , and may be characterized as an extremely elongated member
of that group, in which the beak-sculpture is not quite so fully developed as in
the other species.

Anatomy (Text-fig. 1, p. 455). — The three smallest of my specimens could not
be satisfactorily examined to ascertain the sex, but the gills did not show the mar-

484

MEMOIRS OP THE CARNEGIE MUSEUM.

siijiial structure, so that they may be males. Of the others, three are males, and
one is a gravid female with eggs. The fact that this was collected on July 20 gives
an indication of the breeding season.

Anal opening {a in text-figure) closed above, without forming a supra-anal.
Closed part (s) two to three times as long as the anal; the latter (a) slit-like, short,
shorter than the branchial ojiening. Inner edge of anal practically smooth.
Anal separated from the branchial by a solid bridge (/) formed by the union of
the mantle-margins. Branchial opening (b) a little over twice as long as the anal,
with distinct papilla) on inner edge; mantle-edges not united in front of it. Palpi
(h) subtriangular, the lower margins slightly convex, the posterior margins con-
nected at the base.

Gills (z and o) rather long and moderately wide. The inner (i) wider than
the outer (o) chiefly in front. Outer gill narrowing behind and before, its anterior
end near the highest iioint of the mantle-attachment-line. The inner gill has an
almost straight lower margin, and is only little narrower anteriorly; its anterior
end is immediately behind the ]ialpi. Inner lamina of inner gill entirely connected
with abdominal sac.

Structure of gills normal. In the male both gills have fine, scattered, and
interrupted interlaminar connections, running jiarallel with the gill-filaments,
but without forming complete septa or water-tubes. In the gravid female, the
eggs are contained in the large middle section of the inner gill (i), leaving free
less than one-fourth of the gill at the anterior and at the posterior extremity, and
also leaving free a narrow zone along the margin. This marsupial part has the
interlaminar connections strongly developed, in the shape of interrupted septa,
forming incomplete, intercommunicating water-tubes, filled by the eggs in a dense
mass, not separated into iilacentse. Since no sterile females are at hand, the exact
arrangement of the interlaminar connections in a face view could not be made out.
The charged marsupium is somewhat swollen and distended, so that the inter-
laminar connections have stretched out. The outer gill of the female has the
structure of the gills of the male.

2. Group of Diplodon granosus.

Shell subovate or subtrapezoidal, rather elongated, not distinctly pointed
behind, straight, not distinctly higher in the posterior part, nor oblique. Beak-
sculpture well developed, but fine, and characteristically cut up into numerous
fine nodules or granulations, thus obscuring the radial arrangement. The granular
sculpture often continued a good distance uiion the disk.

ortmann: south American naiades.

485

The essential character of this group is in the beak-sculiiture, which, however,
is very variable, although the granular structure is always more or less evident.
It is hardly possible to give any additional characters, except that the hinge-
teeth are moderately developed, subcompressed, and hardly ever stumjiy. Two
pseudocardinals are in each valve, but sometimes there are reductions.

I have very little material representing this group. The soft jiarts of only
two specimens are at hand, one too young to be of any value, the other a male.
Thus nothing can be said about the structure of the female. All my siiecimens
seem to belong to one species, which is very variable, and has a very limited range
in the coastal streams of eastern Brazil.

The genetic connections of this group likewise cannot be properly ascertained.
The beak-sculpture does not seem to be. very primitive, when compared with the
other species of the geiiUs. In the shape of the shell and the hinge, there is simi-
larity to the next group {chilensis), which ])ossibly may be the most primitive of
the genus.

5. Diplodon granosus (Bruguiere) (1792).

Compare: Unio multistriatus Lea, Von Ihering, 1890, p. 165.

Unio psammactinus Bronn, Von Ihering, 1893, p. 107.

Diplodon expansus Von Ihering, 1910, p. 134,

Diplodon granosus (Brugiere), Simpson, 1914, p. 1250.

I am inclined to accept the earlier opinion of Von Ihering (1890), and that of
Simpson, that a number of so-called “species,” of which Simpson has given a full
synonymy, belong here. Possibly several others should be included.

Type-locality. — Brazil and Guiana.

Additional Localities. — ^Rio de Janeiro (psammactinus, Philippi) (Von Ihering) ;
coastal streams in eastern Brazil, between Rio de Janeiro and Bahia, but not to
the south of Rio (Von Ihering); Rio Negro, Prov. Rio de Janeiro, Distr. St. Rita
(Bunker’s coriaceus, Von Ihering); Tayuara, Distr. Canta Gallo, Prov. Rio de
Janeiro (Von Ihering); Rio Parahyba do Sul, Rio de Janeiro (Von Ihering); Nova
Friburgo, Rio de Janeiro (Von Ihering). Rio Paraguassu, Bahia (Von Ihering,
1910, p. 139, as multistriatus Lea). Rio Doce, Espirito Santo (A^on Ihering, 1910,
p. 134, as expansus Kuester).

New Localities. — Mountain creek, Raiz da Serra, near Santos, Sao Paulo,
Brazil (J. D. Haseman coll., July 26. 1908). One specimen with soft parts, male.

Rio Ribeira, Iporanga, Sao Paulo, Brazil (J. D. Haseman coll., December 1. 1908).
Two specimens, young, one with soft parts of the male type.

486

MEMOIRS OP THE CARNEGIE MUSEUM.

In addition, the Carnegie Museum possesses two specimens labeled “ ellipticus,
Brazil,” from the Holland Collection, and one specimen labeled “ multistriakis,
Brazil,” from the .Tuny Collection.

Dutribution. — The new localities represented in the collection made by Hase-
man extend the range southward (southwestward) along the coast of Brazil beyond
Bio de .Janeiro into the southern portion of the state of Sao Paulo in the small
coastal streams. The estalilished range thus reaches from the Rio Paraguassii
in the North to the Rio Ribeira in the South.

Remarks. — My material is entirely insufficient for the study of the various
forms regarded as belonging here. Originally Von Ihering was inclined to unite all
these forms with granular beak-sculpture into one species; but later he divided them
according to the character of the beak-sciiljiture, although he admits that there is
great variation in this resiiect. Simpson unites them again, at least in part, but
he lets granuliferus Dunker stand, and even adds a new species, D. semigranosus.^^

My specimens also vary in the development of the beak-sculjiture and in the
shape of the shell, but they might very well be forms of one and the same species.
The two soft parts at hand are those of males, and the structure of the female is
unknown.

The two young specimens from Iporanga show the granular beak-sculpture
very well, and I should call them D. grnnosus by all means. 8im])son (1914, p.
1249) has described from the same river-system (Rio Rilieira) at Iguape, Sao Paulo,
Brazil (near the mouth), a D. inirnus, and Marshall (1917, p. 383, pi. 51, figs. 3-6)
has redescribed and figured it. It is founded u])on two specimens, larger than
mine, and differing somewhat from each other in their proportions, and also from
my specimens, but the differences may be individual. The measurements are :

Lengtli.

Height.

Diameter.

Simpson

37 mm.

28 mm.

= 76 pr. ct. of L.

17 mm

= 46 pr. ct. of L.

Marshall

45 “

27 “

= 60

15

= 33

My s])eciinen

29 “

17.5 “

= 60

8.5 “

= 29 “

Do

IS “

11.5 “

= 64

5

= 28 • “

My specimens agree very well with the figures and description, except that there
is no lurid-purple in the nacre, which is dull (lurid) whitish.

Habitats: “Sao Paulo River” (location unknown); “Ponte Grande” (location unknown);
“ Os Perns,” Sao Paulo, Brazil. Marshall (1917, p. 387) redescribes and figures this species, and gives
as type-locality: Rio Tiete, Sao Paulo, and the additional localities “ Ponte Grande, Sao Paulo; Os
Penis; and Ponta Grossa, Parana.” The location of Ponte Grande is still unknown. Ponta Grossa
is on the headwaters of the Rio Tibagy, tributary to the Parand Paneina.

ortmann: south American naiades.

487

The beak-sculpture of Simpson’s species is unknown, being entirely eroded.
It may be that my specimens are this, but I hesitate to unite D. 7nvmus with granosus,
before the beak-sculpture of the former has been described.

3. Group of Diplodon chilensis.

Shell rather compressed, flattened upon the sides, subelliptical, subovate,
subtrapezoidal, more or less elongate, not distinctly pointed behind, straight,
and not distinctly higher in the posterior part, nor oblique. Beak-sculpture
simple, with narrow, straight, uninterrupted radial bars, restricted to the region
of the beaks, and not extending far upon the disk. Two of the median bars may
be joined at their lower ends.

The generally subtrapezoidal shape of the shell, with flat sides, and the simple
character of the beak-sculpture are the chief characters of the group, which may
represent the most primitive type within the genus. The posterior end of the
shell is mostly not pointed, but more or less rounded, and the posterior ridge is
rather indistinct, not prominently angular.

A great number of “species” belong here, which are extremely hard to dis-
tinguish. It is in this group that I encountered the greatest difficulties in the
identification of the species, and even at present I am not satisfied with the results.
This is the more to be regretted, since I have abundant material with soft parts of
some of these forms and have found that there are differences in the anatomy,
which to all appearance are of specific value.

After many attempts to group these forms, I have finally concluded that the
best way, the one that is least liable to lead into error, is to treat these forms geu-
graphically. It is not very likely that the same identical species occurs in widely
separated river-systems.

Forms of this type are found over nearly the whole continent, but apparently
they are rare or missing in the region of the great depression which runs from the
La Plata up the Paraguay to the Amazon-drainage. I shall begin with the forms
from Chile and Patagonia, and proceed then in the direction toward Brazil, going
from South to North,

1. Species from Chile and Patagonia.

I have very insufficient material of this group, and cannot express any positive
opinion about the species. But it seems that all, or nearly all, of the forms known
from Chile belong here. Simpson (1914, p. 1257) admits ten of them: chilensis
Gray, solidulus Philippi, gassiesi Kuester, aplatus Reeve, molince Philipjii, modestus

488

MEMOIRS OF THE CARNEGIE MUSEUM.

Kuester, airatus Sowerb}^, obtusus D’Orbigny, chiloensis Kiiester, aureus Simpson.
Blit the differential characters of these are very obscure, and I should not be as-
tonished if some of these names should prove to be synonyms.

Similar forms are known from the eastern foot of the Cordilleras in Patagonia,
in the drainage of the Rio Negro. Of these the following material is at hand:

6. Diplodon patagonicus (D’Orbigny) (1835).

Unio patagonicus D’Orbigny, 1843, p. 610, PI. 70, figs. 1-4.

Diplodon patagonicus Simpson, 1900, p. 885; 1914, p. 1275; Pilsbry, 1911, p. 610.^^
Type-locality . — Rio Negro, 10-12 miles above its mouth.

New Locality. — Rio Limay, Patagonia (Received in exchange from W. Israel),
one right valve.

Distrihiition. — Rio Negro and its tributary Rio Limay in Patagonia.

The specimen at hand is very poor, with the epidermis worn off, but it is un-
doubtedly this species, which is characterized by its elongated outline and a shallow
radial groove upon the posterior slope.

This species is not at all related to D. parallelopipedon (Lea), with which
it has been associated by Simpson. It has, indeed, a similar elongated outline,
but the characteristic strong and elevated posterior ridge of the latter species is
entirely absent.

7. Diplodon frenzeli (Von Ihering) (1893).

Unio frenzelH Von Ihering, 1893, p. Ill, PI. 4, fig. 12.

Diplodon huapensis Bartsch, 1906, p. 394, PI. 27, fig. 1 ; PI. 28, fig. 1; PI. 29, fig. 2.
Diplodon frenzeli and huapensis, Simpson, 1914, pp. 1264, 1265.

Ty pe-loca lily. — Patagonia.

Other Localities. — Patagonia, foot of the Cordilleras (Von Ihering); small
lake on Victoria Island in Lake Nahuel Huapi, Argentina (Bartsch, huapensis) .

Loccdity Represented in Carnegie Museum. — Lake Nahuel Huapi, Argentina
(C. 11. Eigenmann coll., 1919). One gravid female with soft parts.

Distribution. — Known from Patagonia, at the foot of the Cordilleras in the
region of Lake Nahuel Huapi (Rio Negro drainage). The locality “Chile” given
by Von Ihering is rather vague; and “Os Perus, Sao Paulo, Brazil,” given by
Simpson for frenzeli is probably incorrect.

IMy material is too iioor to give a full account of this species, but I am sure
that my only specimen belongs here, and I also believe that huapensis is the same.
Unio patagonicus Reeve, XVI, 1865, PI. 21, fig. 93, is not this.

ortmann: south American naiades.

489

This species has the indifferent and uncharacteristic outline of the forms of
the chilensis-groxip: subelliptical or subtrapezoidal, with upper and lower margins
subparallel, and rather elongated and compressed shell. It is, however, remarkable
for the anterior location of the beaks (18 or 19 pr. ct. of the length). There is no
radial furrow or groove on the posterior slope.

The color of the epidermis of my specimen (the largest known) is dark brown,
and the epidermis is wrinkled with concentric lamella3, but it has been largely
eroded. According to Von Ihering, the color is dark brown in the larger specimens,
but dark olive in smaller ones, sometimes with lighter green in places. The color
of huapensis is brown posteriorly, grading to wax-yellow anteriorly.

In all other respects, chiefly in the hinge-teeth, my specimen agrees with the
description of frenzeli; only the posterior end of the shell is a little more broadly
rounded, but not very different from the specimen figured by Von Ihering on plate
4, fig. 12i.

D. huapensis has been compared by Bartsch with f renzeli, and he says that it
can readily be distinguished from it by the narrower outline, that means to say by
the height being less in proportion. This, however, is not correct, as can be seen
by comparing the measurements.

The only difference I can see in huapensis is the more tapering posterior end.
But since this species is founded upon two specimens, of which only one has been
figured, this might very well be an individual character.

In the following measurements I leave out Von Ihering’s specimens from Chile,
which appear to me a little doubtful.

Measurements.

Lcngtli.

Height.

Diainetcr.

Beaks.

Huapensis (figure)

53 mm.

25.5 mm

=48 pr. ct. of L.

14

mm. =26 pr. ct. of L.

at 10

mm. =19 pr. ct. of L.

Do. (type, text)

55 “

25.9 “

=47

14.5

“ =26

Do. (text)

57 “

27.3 “

=48

16

“ =28

Frenzeli 5b Von Ihering

56 “

26

=46

16

“ =29

10

“ =18

Do. 5a Von Ihering

58 “

28

=48

17

“ =29

11

" =19

Do. Von Ihering

68 “

34

=50

20

“ =29

= 18

My specimen ( 9 )

71 “

36.5 “

=51

16

“ =23

13

“ =18

Thus my specimen is more compressed than any of the others; however, a
variation of the diameter from 23 to 29 pr. ct. is not at all unusual.

Anatomy. — The specimen at hand has been preserved with the soft parts, and
proved to be a gravid female with glochidia.

Color of soft parts whitish, with black pigment near anal and branchial open-
ings extending forward a good distance along the margin of the mantle, and be-
coming brown.

490

MEMOIRS OF THE CARNEGIE MUSEUM.

Bartsch has given a description of the soft parts of huapensis, which refers
chiefly to the general features, the color, the structure of the siphons, and the
shape of the gills and palpi. But no particulars as to the structure of the gills are
given.

Anal opening slit-like, closed above, closed part about four times as long as
the open, the latter with the inner edge smooth, and separated from the branchial
by a solid mantle-connection. Branchial opening with distinct papillae. Palpi
of moderate size, subtriangular, lower margins curved, posterior margins not
connected at base.

Gills nearly of the same width posteriorly, the inner much wider than the
outer anteriorly. Outer gill subtriangular, its anterior end at the highest point
of the mantle-attachment-line. Inner gill with gently convex margin, very little
narrower anteriorly, broadly attached in front, its anterior end immediately be-
hind the palpi. Inner lamina of inner gill entirely connected with abdominal sac.
Interlaminar connections of non-marsupial gills weakly developed, scattered.
Marsnphivi located in the inner gill, but not occupying all of it, leaving nearly
one-fifth of the gill free anteriorly, and about two-fifths posteriorly, so that the
marsupium is distinctly shifted forward, lying in the second and third fifth of the
gill (from the anterior end). Interlaminar connections of the marsupium forming
interruiited sejita, the septiform structure apparently prevailing (this is not quite
clear, since the structure is obscured by the masses of glochidia). The charged
marsupium is a little swollen, and the glochidia fill it in a loose mass, not being
conglutinated.

Glochidimn subtriangular, strongly obliciue, with the point of the lower margin
located vertically below the posterior end of the hinge-line. Hooks are present
and of the normal 8-shape. L. 0.25, H. 0.20; L. of hooks: 0.05 mm. Compared
with other species, the glochidium is rather small.

2. Species prom coastal streams of southern Brazil.

It is a noteworthy fact that species of the c/u7e/isis-type seem to be absent
from the system of the Rio de la Plata (with the exception of the drainage of the
upper Parana). They also seem to be absent in the great Amazons-basin, and
northward. But they are found rather plentifully on the Brazilian plateau, be-
ginning at its southern extremity, in the coastal streams in Rio Grande do Sul,
going thence northward, and crossing over in Sao Paulo and Parana into the drain-
age of the iipiier Parana River.

We shall take up first the species of the coastal streams, and I wish to call

ortmann: south American naiades.

491

attention (as has been done by Simpson in the case of D. mimus) to the great re-
semblance of these shells, not only among themselves, but also to the North
American Elliptio complanatus (Dillwyn), distributed over the Atlantic streams
of the eastern coast of the United States from Georgia to Maine. This resemblance,
of course, is only external and superficial. Closer examination of the hinge, of
the adductor-scars, and, if visible, of the beak-sculpture, at once reveals important
differences. The anatomy is entirely different.

8. Diplodon imitator Ortmann, sp. nov.

Shells: PI. XXXIV, figs. 5, 6, 7; PI. XXXV, figs. 1, 2. Anatomy of gills: PI. XLV,
fig. 1. Sectio7i of gills: PI. XLVII, fig. 6. Glochidium : text-fig. 46 (p. 469).

Type-locality. — Rio Vaccahy-mirim, Santa Maria, Rio Grande do Sul, Brazil
(J. D. Haseman coll., January 29. 1909). Twenty-three specimens, all with

soft parts, among them males, barren and gravid females, with eggs and with
glochidia. Type-set: Cam. Mus. Cat. No. 61.9248. (This is in the drainage of
the Rio Guahyba-Jacuhy, far up in the headwaters).

Additional Locality. — Rio Jacuhy, Cachoeira, Rio Grande do Sul, Brazil (J. D.
Haseman coll., January 26. 1909). One barren and one gravid female, with eggs,

both young. (This is farther down, in the middle part of the Rio Jacuhy.)

(Originally there were thirty-four specimens from the type-locality at my
disposal, all with soft parts.)

Only once before has a form of this group been reported from the Guahyba-
drainage, viz. D. martensi Von Ihering, from Taiiuary and Santa Cruz (probably
from tributaries of the lower Jacuhy (See Von Ihering, 1893, p. 102). . But these are
not typical martensi, the original of the latter being ‘‘probably from Sao Paulo).
The dimensions of these specimens from Rio Grande do Sul given by Von Ihering
agree fairly well with those of the real U . martensi, chiefly the relation of height
to length: 49 pr. ct. in two specimens from Rio Grande do Sul, 49 pr. ct. (text) or
53 pr. ct. (figure) in martensi. But these dimensions do not agree with my speci-
mens, where the height ranges from 55 to 64 pr. ct. and never falls as low as in
martensi. The diameter of martensi is also greater than in my material. For
this reason I am comiielled to describe my shells as a new species.

Description of Shell. — Of medium size (maximum length 80 mm.), moderately
solid, rather thin when young, compressed (diameter 26 to 33 pr. ct. of length),
subelliptical, subovate, or subtrapezoidal, moderately elongated (height 55 to

It is much to be regretted that the type-locality is not better known. It is very doubtful, on
account of the great similarity of these forms, whether the real martensi can ever be positively identified.

492

MEMOIRS OP THE CARNEGIE MUSEUM.

64 pr. ct. of length). Valves not gaping. Dorsal margin nearly straight or very
gently convex, passing gradnall}^ into the anterior margin (rarely forming an in-
distinct angle). Posterior upper margin forming a blunt angle with the posterior
margin or passing gradually into it. Posterior margin obliquely descending,
more or less curved, and curving more strongly into the lower margin, without
forming a distinct posterior angle. The lower posterior end of the shell is rounded,
but hardly biangular. Lower margin in young specimens gently convex, in older
ones it is rather straight in the middle. Anterior end of shell not, or very little,
narrower than the posterior end. The shell is thus rather straight and not oblique.

Valves only slightly convex, rather flattened upon the sides, with the posterior
ridge very indistinct and broadly rounded. Posterior slope somewhat compressed.
Greatest diameter near the middle of the shell, but not posterior to it. Beaks
not swollen, and not elevated, located at from 20 to 28 pr. ct. of the length. Beak-
sculpture consisting of about fourteen to sixteen rather sharp and fine, short, radial
bars, 5 to 7 mm. long, those upon the posterior ridge hardly longer than the rest,
and none of them uniting in the middle of the shell with their lower ends. No
distinct granulations present, but sometimes there are a few irregular oblique
wrinkles upon the posterior slope near the beaks. A short, narrow lunula in older
shells.

Epidermis in young specimens rather smooth and shining, but with fine, ir-
regular, concentric striie, nowhere lamellar. In old specimens it is less smooth,
chiefly on the iiosterior slope and toward the lower margin with more crowded
and rougher striae. ( I'inkled radial lines hardly indicated upon the shell. Color
in young specimens greenish bronze or brownish, sometimes with indistinct brown-
ish concentric bands, in older shells greenish tints disappear, and the epidermis
is dull brown or blackish brown.

Hinge-line gently curved. Ligamental sinus over the middle of the lateral
tooth or slightly behind it, in older shells over its last third. Laterals curved, one
in right, two in left valve, somewhat rough posteriorly. Pseudocardinals normally
two in each valve. In young specimens those of the right valve are obliquely and
forwardly descending, compressed, the anterior low and narrow, the posterior
higher, and a little thicker, crenulated. In older specimens the posterior is thicker
and becomes generally more triangular and stumpy. Those of the left valve of
young specimens are also compressed, but subtriangular and not very long, crenu-
lated, the anterior one larger than the posterior. In old shells these two teeth are
more stumiiy, triangular, and not compressed. In rare cases reductions take place,
chiefly with regard to the anterior tooth of the right valve, or with regard to the

ortmann: south American naiades.

493

posterior in the left valve, which may become very small, and sometimes an ac-
cessory third posterior tooth may develop in the right valve. Thus the pseudo-
cardinals are quite variable in number, shape, size, compression, and development
of rugosities.

Cavities of shell and beaks shallow. Nacre blueish white or white, often
discolored and with lurid tints (grayish purple) toward the beak-cavities, iridescent
posteriorly.

Anterior adductor-scar distinct and impressed, subtriangular. Anterior
retractor-scar separated from it, small, round, deeply impressed. Anterior pro-
tractor-scar connected with adductor-scar, forming a posterior process of it. Pos-
terior adductor-scar shallow, subtriangular, with an upper process formed by the
posterior retractor-scar. Pallial line distinct. Dorsal muscle scars a few, lying
in a line in the beak-cavity.

IMeasurements.

No.

Sex.

Length.

Height

Diameter.

Beaks.

Figure.

11.. .

cf

40.5 mm.

24 mm. =59 pr. ct. of L.

12

mm. =30 pr. ct. of L.

at 11

mm. =27 pr. ct. of L.

16.. .

d'

43

27 “ =63

11

“ =26

11

“ =26

21.. .

9

47

30 “ =64

15

“ =32

11

“ =23

PI. XXXV, fifr. 2.

29.. .

d

62

34 “ =55

20.5

“ =.33

14

“ =23

PI. XXXIV, fig. 5.

32.. .

33.. .

9

9

76

80

44.5 “ =59

44 “ =55

23.5

26.5

“ =31
“ =33

16.5

16

“ —22 “

“ =20

f PI. XXXIV, fig. 7.

1 PI. XXXV, fig. 1.

Anatomy. — Anal opening slit-like, shorter than branchial opening, closed
above; closed part about twice as long as open jiart, no supra-anal formed. Bran-
chial opening separated from anal by a solid connection of the inner mantle-edges.
Inner edge of anal almost smooth, that of branchial with distinct papillae. In
front of the branchial the mantle-edges are unconnected. Palpi subtriangular,
a little longer than wide, their posterior margins connected at base.

Gills long, the outer subtriangular, wider in the middle than the inner. The
inner not triangular, wider than the outer anteriorly, its anterior end attached
to the space between anterior end of outer gill and palpi, and in contact with the
posterior base of the latter. Inner lamina of inner gill entirely connected with
abdominal sac.

Non-marsupial gills (PI. XLV, fig. la) with few, scattered, irregularly dis-
posed interlaminar connections. Marsupium (PI. XLV, fig. 16) located in the
inner gills, occupying only a part of them, at and in front of the middle; at anterior
end about one-fifth of the gill remains non-marsupial, and at the posterior end
about two-fifths, so that the marsupium occupies two-fifths of the gill, the second
and third from the anterior end. This location of the marsupium is constant for

494

MEMOIRS OF THE CARNEGIE MUSEUM.

the species (observed in seventeen females), with the only cpialification that in
young females the marsupium is smaller (less than two-fifths of the gill) but has
a similar position.

Marsupial part formed by interrupted septa (PL XLV, fig. 16; PL XLVII,
fig. 6). The arrangement into septa is distinct, but they are frequently inter-
rupted, and their solid portions are short. In the middle of the marsupium they
are at the utmost six to eight times as long as thick, and in its other parts they are
very short. A distinct (luincuncial (reticulate) arrangement is not seen. Toward
the margin of the gill the septiform arrangement is again more distinct.

The embryos fill the marsupium in an irregular mass, and the charged mar-
supium is moderately swollen. Placentie are not formed.

Glochidiuni (text-fig. 46, p. 4G9, observed in six specimens) subtriangular,
slightly obli(iue, anterior and posterior margins convex, converging to a point,
anterior margin longer than the posterior. Hooks present, of the Hyriine type, with
the S-shaped curve. L. 0.27 to 0.28 mm. ; H. 0.27 to 0.28 mm. ; L. of hook: 0.09 mm.

It should lie remarked that in one female the glochidia were immature, and
no hooks could be seen : three females had only eggs, and one female was in the act
of discharging. The dates of collecting, Jan. 20 and 29, should be recorded as
showing the breeding season (midsummer of Southern Hemisphere).

Remarks on the Specific Characters. — As will be seen from the descriptions
of the following species, D. sirnillimus, D. vicarius, and D. decipiens are very closely
allied to the present species in the shape of the. shell; vicarius can be readily dis-
tinguished by the biangular posterior ridge, and D. sirnillimus is a smaller shell.
In other respects it is almost impossible to distinguish these species by the shell
alone. The few obscure, differentiating characters will be pointed out below under
the resiiective species. However, the investigation of the soft parts has shown
that there are interesting differences chiefly in the location of the marsupium. In
the present species, the marsupium is most fully developed, occupying a rather
large part of the inner gills, slightly gravitating toward the anterior end. In the
following species (chiefly simillimus and vicarius) it will be seen that this tendency
is increased, and the marsupium becomes smaller, and is being shifted more dis-
tinctly forwards (Compare pi. XLV, fig. 16, with. PL XLV, figs. 26, and 3). It
will also be seen that there are slight differences in the glochidia with regard to
obliquity and size.

ortmann: south American naiades.

495

9. Diplodon simillimus Ortmann, sp. nov.

Shells: PL XXXV, figs. 3, 4, 5, 6. Anatomy of gills: PI. XLV, fig. 2.

GlocMdium: Text-fig. 4c, p. 469.

Type-locality. — Rio Nhimdiaquara, Morretes, Parana, Brazil (J. D. Haseman
coll., January 3, 1909). Type-set: Cam. Miis. Cat., No. 61.9250. Thirty-two
specimens with soft parts, males, barren and gravid females with glochidia.

About a dozen additional specimens belonging to the same original lot have
been studied. The locality is in a small coastal stream emptying into the Bay of
Paranagua.

No shells have ever been reported from the region of Paranagua Bay in Parana.
However, from a little less than one hundred miles to the south, in Santa Catharina,
near Barra Itapocu (I believe that the Rio Itapoca, as given by Marshall, stands for
this), Diplodon santamarice Simpson has been described (Simpson, 1914, p. 1270;
and Marshall, 1917, p. 386, PI. 52, fig. 6; PI. 55, figs 1-4). This is founded upon
three specimens only, and resembles our species to a degree. But judging from
description and figures, it is somewhat larger (max. 63 mm.), longer (H. 52-59
pr. ct. of L.), and the hinge has the posterior tooth of the left valve missing. Since
nothing is known of the anatomy of D. santamarice, and the locality is not the same,
it would be rash to unite our specimens with this sjiecies.

Our species also is much like U. martensi Von Ihering, and might fall under
this according to Von Ihering’s conception. But it cannot be united with it on
account of the different dimensions. While in martensi the height is said to be
from 49 to 57 ])r. ct., our specimens are mostly less elongated, with the height from
53 to 66 pr. ct. of the length. In D. martensi the diameter is 32 to 35 pr. ct., in the
present species from 26 to 37 pr. ct. (this agreeing better with martensi than with
imitator). But in the absence of exact localities for martensi and any knowledge
of its anatomy, and in view of the general resemblance of all of these shells, it is
impossible to identify our shell with any previously described, and no other alterna-
tive exists, except to describe it as new.

In the characters of the shell D. simillimus is very close to D. imitator. The
description of the latter species would fit it very well, and I shall here only emphasize
the distinguishing characters.

1. D. simillimus is a smaller shell (max. length 61 mm., as against 80 mm. in
imitator) .

2. The trapezoidal shape, with an angle between the upper and the posterior
margins, is seen here only in very young shells. In older shells these two margins
form a rather regular curve.

496

MEMOIRS OF THE CARNEGIE MUSEUM.

3. The lower margin of the shell of D. svmillimus is frequently more nearly
straight, so that the shell of older specimens a]ipears more humped.

4. Color of epidermis with hardly any green tints, but light to dark bronze-
brown, in old shells brown-lilack.

5. Pseudocardinal teeth never stumpy, chiefly in left valve, but always com-
pressed, although the two of the left valve and the ]iosterior in the right are rather
thick in old specimens.

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Figured.

5.. .

c?

SG.-I mm.

21.5 mm

= .59 pr. et.. of L.

9.5

mm. =26 pr. r

t. of L.

at, 8.5

mm. =23 pr. ct. of L.

29.. .

cf

37.5 “

23

= 61

11

“ =29

“

10

“ =27

18.. .

9

42.5 “

22.5 “

= .53

11

“ =26

“

9.5

“ =22 “

11.. .

&

45

27

= 60

13

“ =29

“

9

“ =20

PI. XXXV, fig.

3.

23.. .

9

59

39

= 66

22

“ =37

“

14

“ =24

22.

9

61

37

= 61

20.5

“ =.34

15

“ =25

PI. XXXV, fig.

5.

The characters of the shells of this species are rather poorly marked, and can
be ascertained only by the examination of extensive material. I have discovered,
however, that there are imiiortant and constant differences in the anatomy.

Anatomy. — Fidly agreeing with that of D. imitator, but the marsupium is
different (See PI. XLV, fig. 2). It is located in the anterior part of the inner gill,
entirely anterior to its middle, and extending forward to within a short distance
of the anterior end of the gill, so that anteriorly less than one-tenth of the gill is
non-marsupial, while posteriorly fully the posterior half of it is non-marsupial.
The interlaminar connections form interrupted sejita, but the septiform structure
is less distinct than in D. imitator, and more of a reticulated (or irregular quin-
cuncial) arrangement is evident. This structure of the marsupium is the same in
all females investigated (altogether twenty individuals), and only in young ones is
the marsupial ]iart smaller.

The Glochidium (Text-fig. 4c, p. 469) differs from that of D. imitator in being
more distinctly oblique, and being longer than high. L. 0.28 mm., H. 0.24 mm.
There are hooks of the same type, which arc about 0.10 mm. long.

My specimens were collected on January 3, which date indicates the breeding
season, jirobably its middle, for eight females had only eggs, while nine had glochidia,
in part not mature, and with the hooks yet unformed.

The remarkable fact lirought out by the study of the anatomy is that, while
the species is extremely hard to distinguish fro7n imitator by the shell, it has at least
two anatomical characters {'marsupium and glochidium) , which are very well ^narked
and constant.

ortmann: south American naiades.

497

10. Diplodon vicarius Ortmann, sp. nov.

Shells: PI. XXXV, figs. 7, 8; PI. XXXVI, figs. 1, 2. Anatomy of gills: PI. XLV,

fig. 3. Glochidium: Text-fig. 4d, p. 469.

Type-locality. — In creeks. Aqua Qiiente (eight miles from Iporanga), Sao
Paulo, Brazil, tributaries of Rio Ribeira (J. D. Haseman coll., November 27, 1908).
Type-set: Cam. Mus., Cat. No. 61.9251; fifteen specimens all with soft parts,
males, barren and gravid females.

Additional Locality. — Rio Ribeira, Iporanga, Sao Paulo, Brazil (J. D. Haseman
coll., December 1. 1908). One specimen, male, with soft parts.

Only one species of Diplodon has been described from the Rio Ribeira; this is
D. mimus Simpson (1914, p. 1249; Marshall, 1917, p. 383, PI. 51, fig. 3) from Iguape,
Sao Paulo, at the mouth of the river. As has been pointed out above (p. 486),
this species might possibl^^ be D. granosus. But on the other hand a few ]3articulars
agree with the present species, as, for instance, the biangulation of the posterior
end and the general resemblance to Elliptio complanatus mentioned by Simjison.
Yet our shells cannot be this, because they are larger, less convex, and have dif-
ferent dimensions. D. niinius is smaller (max. 45 mm.), and has, according to
the measurements given, a considerably higher (60 to 76 pr. ct.) and more swollen
(D. 33 to 46 pr. ct.) shell, while D. vicarius has the height only from 52 to 63 pr. ct.
and the diameter from 26 to 31 jir. ct., and is a good deal larger (L. 53 to 68 mm.).
Of course, our form may fall under martensi Von Ihering, but for the same reasons
as in the case of the two preceding sjiecies, it cannot be called by this name, and
thus we must describe it as new.

It may perhaps be that D. vicarius, of which we do not know the beak-sculp-
ture, is the older stage of the young s})ecimens recorded from Iporanga as D.
granosus (See p. 485). The shape and dimensions agree fairly well, but the size
is very different, the maximum length of granosus lieing only 29 mm. and no inter-
mediate specimens between these and minimum length of vicarius (53 mm.) are
at hand. Thus the question must remain unsettled. D. granosus from other
localities is also always much smaller than vicarius.

This species is also extremely similar to D. imitator and D. simillimus. In size
it stands between them (max. length 68 mm.). The outline is also subtrapezoidal
or subelliptical, and the shell is quite compressed, resembling the shape of Elliptio
complanatus of the United States. The lower margin in the shells before me is
always rather straight, but I have not very young specimens. However, from the
growth-lines it is seen that young shells must have had a gently curved lower margin.
In none of my specimens is the beak-sculpture preserved.

498

MEMOIRS OF THE CARNEGIE MUSEUM.

Characters of the Shell. — The description of D. imitator might also serve for
this species. However, the following differences are noticeable:

1. Posterior ridge of shell broad, and more or less distinctly biangulate, pro-
ducing a biangiilation also of the posterior end of the shell. This is the most
striking character of the shell. In both D. imitator and simillim.us the posterior
end of the shell is evenly rounded without any trace of angulations.

2. Epidermis not so shining as in the two other species, which is due to the
development of additional fine concentric wrinkles, which are irregular and best
developed ujion the posterior slope and near the lower margin. Upon the disk
they form indistinct radiating lines (or narrow bands), which are obsolete in the
two other species. Color of epidermis of vicarius lighter or darker brown, without
gi’een tints, and without bronzy lustre.

The hinge-teeth are generally of the type of D. simillimus, i.e., they do not
become stumpy in old shells. They are, however, very variable, and much cut
up, more so than in the two other species, and the posterior pseudocardinal of
the left valve frecpiently is quite small or rudimentar}^ The nacre is whitish, but
often inclines to lurid tints (purplish gray).

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Figured.

13.. .

d’

53 nim.

31 mm. =58 pr. ct. of L.

15.5 mm. =29 pr. et. of L.

at 12 mm. =23 pr. ct. of L.

PI. XXXV, fig. 7.

9.. .

9

53.5 “

33.5 “ =63

14 “ =26

12 “ =22

PI. XXXVI, fig. 2.

10.. .

9

.57

34.5 “ =61

16 “ =28

11.5 “ =20

1.. .

9

58.5 “

33.5 “ =.57

15 “ =25

1.5 “ =26

G.. .

(d

G7

.3.5 “ =52

19 “ =28

16 “ =24

.

15.. .

9

68 “

40 “ =.59

21 “ =31

15 “ =22

) PI. XXXV, fig. 8.
iPl. XXXVI, fig. 1.

No sexual differences in the shell.

Anatoimj. — I have eight females, six of which are gravid; three with eggs, three
with glochidia (immature in one). The structure of the soft parts is exactly as in
the preceding species, except that of the marsupium (Plate XLV, fig. 3). As in
D. simillimus, the marsuiiium is located in the anterior portion of the inner gill,
anterior to the middle, but it occupies a still smaller part, and does not extend so
near to the anterior end, and does not reach the middle of the gill. Anteriorly
about one-ninth of the gill is non-marsupial, and posteriorly over half of it. The
interlaminar connections are arranged in interrupted septa, the septiform arrange-
ment being most evident anteriorly and in the middle of the marsupium, while
posteriorly the arrangement is reticulate (indistinctly quincimcial).

The glochidium (Text-fig. 4d, p. 469) is similar to that of D. simillimus, oblique,
but slightly longer. L. 0.30, H. 0.24 mm. There are hooks of the same type, which

ortmann: south American naiades.

499

are at least 0.09 mm. long. In one specimen with immature glochidia no hooks
could be seen.

In this case also the breeding season falls in the winter months of the northern
hemisphere, but apparently a little earlier than in the other species (end of No-
vember).

Thus this species has anatomical characters of its own, chiefly observable in
the size and location of the marsupium. It agrees most closely with D. simillimus.

3. Species from the drainage of the upper Parana.

I have material belonging to the c/u7ensfs-group from the following tributaries
of the Parana: Rio Iguassu, Rio Tiete, and Rio Grande in Sao Paulo. A form from
the Iguassu is noticeably very closely related to the three species from the coastal
streams just described. We should bear in mind that the head-waters of the
Iguassu, from which this form comes, are in close proximity to those of the Rio
Nhundiaquara and Rio Ribeira on the eastern watershed.

11. Diplodon dectpiens Ortmann, sp. nov.

Shells: PI. XXXVI, figs. 3, 4, 5, 6. Anatomy of gills: PI. XLV, fig. 4. Section of
gills: PI. XLVII, fig. 7. Glochidium: Text-fig. 4c, p. 469.

Type-locality. — Creek, tributary to the Rio Iguassu, Serrinha, Parana, Brazil
(J. D. Haseman coll., December 23. 1908). Type-set: Cam. Mus. Cat. No.

61.9253, thirteen specimens, males, barren and gravid females, all with soft
parts.

No Naiades have hitherto been known from the river-system of the Iguassu,
and, as in the preceding species, none of the names of species which may occur here,
can be applied to our specimens with any degree of certainty. Therefore we in-
troduce this form as a new species.

In the shape of the shell this species is very close to the three preceding, es-
pecially imitator and simillimus , which it resembles in its subelliptical or sub-
trapezoidal outline. The latter shape is seen chiefly in younger specimens, while
older ones become more subelliptical. The posterior ridge and the posterior end
are never biangulate as in vicavius. In the glossy epidermis, D. decipiens also
agrees better with imitator and simillimus, and the radial lines formed of fine
wrinkles are poorly developed. The general description given for imitator might
be repeated for this species, and the beak-sculpture in particular has the same
character. Nevertheless the following peculiarities should be mentioned as
diagnostic :

500

MEMOIRS OF THE CARNEGIE MUSEUM.

1. The beak-sculpture has a smaller number (ten to twelve) of radial bars,
the bars having the same length (5 to 7 mm.)

2. The outline of the shell is more freipiently subelliptical, often rather regularly
so, with both upper and lower margins almost equality curved. In some specimens,
indeed, the lower margin is more nearly straight, but this never causes a distinctly
hum])ed shape (so often seen in simillvm.us) . No trace of biangulation posteriorly,

3. Epidermis in young shells bronzy-brown or bronzy-green; in older ones it
becoming a deep chestnut-color, inclining jiartly to blackish.

In size this sjiecies stands between imitator and vicarius (maximum length
73 mm.). The relative dimensions are very much like those of the other three
species. The hinge-teeth most resemble those of simillimus, never being stumpy,
as in imitator] they are not much cut up; and the iiosterior pseudocardinal in the
left valve has a strong tendency to disapjiear, being sometimes entirely missing.
The nacre is whitish, l)ut generally lurid (iiuriilish gi’ay) in the cavity.

IMeasurements.

No.

Sex.

I.ength.

Ileiglit,.

Diameter.

Beak.s.

Figured.

a. . .

d’

26 nun.

14.5 inin. =50 pr. ct,. of L.

0 mm. =23 pr. ot. of I...

at 0.5 mm. =25 pr. ct. of L.

c, . .

9

35 “

20.5 " =59

9.5 “ =27

7.5 “ =21

PI. XXXVI, fig. 6.

. .

9

58 “

32.5 “ =50

18.5 “ =32

15 “ =26

PI. XXXVI, fig. 5.

4.. .

d'

07 “

30.5 “ =54

20.5 “ =31

15 “ =22

PL XXXV, fig. 3.

10.. .

9

70 “

38 “ =54

25 “ =30

10 “ =23

0.. .

9

73 “

40 “ =.55

25 “ =34

10 “ =22

PI. XXXVI, fig. 4.

No sexual differences in the shell.

Anatomy. — Aside from three very young specimens, which probably are a male
and two females, I have six males of good size, and one barren female, a gravid
female with eggs, and two females with glochidia (in one of them immature).

The anatomy is similar to that of D. imitator, smiillimus, and vicarius, but
the marsupium (PI. XLV, fig. 46) is quite different, essentially differing from that of
the other species. It is located in the middle of the inner gill, leaving between one-
fifth and one-fourth of the gill at the anterior end, and about one-third of it at the
posterior end non-marsiq)ial. Thus the marsiqiial portion is rather large, and more
of it is anterior rather than jiosterior to the middle. This location comes nearer
to what is seen in D. imitator, than in the other two species. The most striking
character, however, which we have not before encountered, is that the interlaminar
connections of the marsupial part are not interrupted, but form continuous septa,
running from near the base of the gill close to the margin. These septa are heavy
and stand very closely, forming regular, isolated water-tubes (See section on PI.
XLVn, fig. 7). The whole of the marsupium has this structure, and no inter-

ortmann: south American naiades.

501

rupted septa are seen anywhere. That these septa are only a modification of the
scattered interlaminar connections is shown by the fact that the latter are found
in the non-marsupial gills (See outer gill on PI. XLV, fig. 45, and gills of male,
PI. XLV, fig. 4a). All four of my large females show this arrangement, and in
the two young ones traces of the beginning of this structure are seen. The em-
bryos fill the water-tubes in loose masses, not forming distinct placentae.

Glochidium (Text-fig. 4e, p. 469) much like that of D. vicarius, oblique, L.
0.31, D. 24 mm., with hooks, 0.09 to 0.11 mm. long. One specimen has immature
glochidia, but rudimentary hooks can be seen. The date of collecting (December
23) gives a hint as to the breeding season.

While this species is ver}^ close to the three preceding in the characters of
the shell, it has differences in the soft parts, which are very striking. The structure
of the marsLipium is extremely interesting, and there is no doubt that it must be
regarded at least as of specific value, rejiresenting a high specialisation of this
organ.

12. Diplodon paulista (Von Ihering) (1893).

Anatomy of gills: PI. XLVI, fig. 1. Section of gills: PI. XLVTII, fig. 1 ;

Glochidium: Text-fig. 4 f. p. 469.

Unio paulista Von Ihering, 1893, p. 93, PI. 4, fig. 7.

Diplodon paulista Simpson, 1900, p. 873; 1914, p. 1229.

Type-locality. — Piracicaba, Sao Paulo, Brazil; according to Nehring (1893, p.
166) in Rio Piracicaba Mirim.

New Localities. — Rio Tiete, Mogy das Cruzes, Sao Paulo, Brazil (Headwaters
of R. Tiete) (J. D. Haseman coll., July 19, 1908) males, barren and gravid females,
originally twent3^-five in the lot, all with soft parts. Rio Tiete, Sapina, Sao Paulo
(Exact location unknown, but must be near city of Sao Paulo) (J. D. Haseman
coll., July 23, 1908) one male with soft parts.- Creek, tributary to Rio Mog^"
Guassu, Mogy Mirim, Sao Paulo, Brazil (tributary to Rio Grande and Parana)
(J. D. Haseman coll., August 7, 1908) males and gravid females, originallj^ seven-
teen specimens in the lot, all with soft parts.

A detailed description of five specimens has been given by Von Ihering, but
the specific characters have remained obscure. The specimens before me agree
very well with this description, but it should be noted that the two forms of the
shell supposed by Von Ihering to belong to the male and female, do not represent
sexual differentiation, but simply individual variations. The more regularly
ovate outline, believed to belong to the male, is in fact rather rarelj" well-developed.

502

MEMOIRS OF THE CARNEGIE MUSEUM.

while the other, somewhat more oblique, is more frequent; but both pass insensibly
into each other. According to my material younger specimens are more apt to
exhibit the more regularly ovate outline.

This species is not very closely allied to those mentioned on the preceding
pages, but represents a somewhat different type, and inclines towards the ellipticus-
groiqi by its often slightly oblique shell. It is much smaller than imitator, vicarius,
and decipiens, and also does not attain the size of simillimus. The largest specimen
before Von Ihering was 57 mm. long, while my largest falls even short of this (45
mm. from Rio Tiete, 51 mm. from Mogy Mirim). The subtrapezoidal shape is only
distinct in very young individuals; in larger specimens it becomes subelliptical or
subovate, generally a little higher posteriorly and slightly oblique. The ground-
color of the epidermis is greenish-olive, and never distinctly brownish, although old
specimens may become blackish green. The beak-sculpture consists of fine, sharp,
and short radial bars, the numlier of which is sixteen to eighteen, the median pair
having a tendency to unite at the lower ends.

The characters of the inside of the shell have been well described by Von
Ihering. The left valve has generally only one jiseudocardinal, but sometimes
there is a trace of a second posterior one. It should be noted that, as described
by Von Ihering, the posterior retractor-scar is separated from, and stands above
the })osterior adductor-scar, but not always, as in some cases it is connected with
it, and this maj" be different, even in the right and left valve of the same individual.

Measurements.

No.

Sex.

Length.

lleiglit.

Diameter.

Beaks.

Von Iherii

ig

48 to 57nun.

59 to 65 pr. et. of L.

31 to 35 pr. ct. of L.

22 to 26 pr.

ct. of L.

M o(]y das Cruzes

?

15. .5 mm.

8.5

min.

=55

4

mm. =20

at 4

mm. =20

18

9

27

10

= 59

8

“ =30

0

“ =22

10

d’

38

22

= 58

11.5

“ =30

7.5

“ =20

2.3

9

38

24

= 63

14

“ =37

10

“ =26

7

cf

41

27.5

= 67

14

“ =34

10

“ =24

14

d'

45

27

= 60

16

“ =36

11

“ =24

Mood Mirim

9

cf

35

22

= 03

14.5

“ =41

10

“ =29

14

d’

39

23.5

= 60

13

“ =33

9.5

“ =24

13

cf

40.5 “

24

= 58

15

“ =37

11.5

“ =28

12

o'

42

28

= 67

18

“ =43

11.5

“ =27

3

9

48.5 “

29

= 00

16.5

“ =34

13

“ =27

5

(T

51

32

= 03

18

“ =35

13

“ =26

Vly two sets from the Rio Tiete and from a creek tributary to Rio Mogy
Cluassu, differ slightly from each other. In the former the size is a little smaller^
and the young shells are rather more elongated, thus rendering the average height
less. The diameter is also not so great in the shells from the Tiete. The measure-

ortmann: south American naiades.

503

ments of the dimensions of the two sets largely overlap, and they agree very well
with those given by Von Ihering. Of course, my material being more plentiful,
the range of the dimensions is wider.

An apparently allied form is D. suppositus Simpson (1914, p. 1245; Marshall,
1917, p. 385, PI. 51, fig. 2; PI. 54, fig. 1-4).^® The type is, according to Marshall,
from ^'Parana, Brazil,” and other specimens are from Rio Tiete, Sao Paulo, and
other localities in southern Brazil. This is also a comparatively small form, but,
according to the measurements given, it is more elongated, the height being in
the type 52 pr. ct. of the length (53 pr. ct. according to Marshall), and in another
specimen 58 pr. ct. This differs somewhat from our specimens (58 to 67 pr. ct.)
and from Von Ihering’s paulista (59 to 65 pr. ct). In addition suppositus has a
chestnut-bronzy epidermis, while it is greenish in paulista. Thus the two forms do
not agree.

Anatomy. — I have investigated the soft parts of all of my shells, and there
were altogether, aside from several small ones, where the sex could not be ascer-
tained, nineteen males, six barren females, five gravid females with eggs, and seven
gravid females with glochidia.

The anatomy in general is like that of the genus Diplodon. But in this case
again the marsupium (PI. XLVI, fig. 1) in its size and location shows specific peculi-
arities, exhibited by all of my females. It agrees with the preceding species in
the fact that it occupies only a part of the inner gill, and in the slightly anterior
location. But the marsupial part is very small, occupying about one-fifth or one-
fourth of the length of the gill, leaving a considerable portion non-marsupial at
the anterior end, and half or nearly half at the posterior end. The figure on PI.
XLVI, fig. 1, represents a specimen with a rather small marsupium; generally it
is a little larger. Thus the marsupium is in the middle of the gill, and slightly
in advance of the middle. When charged, it forms here a rounded or oval swelling,
rather distant from the base, and extending not quite to the edge. There is a
slight variation in the specimens from the two main localities. In those from Mogy
das Cruzes it is distinctly in front of the middle; in those from Mogy Mirim more
median, but this difference is very slight and not always distinct. The inter-
laminar connections form interrupted septa (also seen in the section, PL XLVIII,
fig. I), and there is no distinctly reticulate arrangement.

Glochidium (Text-fig. 4/, p. 469) rather large, subtriangular, oblique, with
hooks. L. 0.32, H. 0.27 mm.; hooks about 0.10 mm. long. (Thus the glochidium
is slightly larger than in any of the preceding species.)

The name suppositus was first given by Von Ihering (1893, p. 102) without description, from
Rio Grande (Upper Parand, boundary between Sao Paulo and Minas Geraes).

504

MEMOIRS OF THE CARNEGIE MUSEUM.

4. Group of Diplodon charruanus.

Shell not compressed, but rather evenly convex and often considerably swollen
when old. Outline subelliptical or somewhat sulitrapezoidal, elongate, more
or less pointed behind, straight, and not oblique. Beak-sculpture simple, with
narrow, straight, uninterrupted bars, restricted to the region of the beaks. A
few of these bars may lie joined at their lower ends.

The chief character by which this group is distinguished from that of chilensis
is the absence of a flattening of the valves upon the sides. The valves are generally
evenly convex, and in consequence of this, at least in larger specimens, the shell
appears as more swollen. In other respects, the shell is similar in outline and other
characters, except that the posterior end is often produced into a distinct blunt
point, which may be more or less elevated aliove the base-line. There is no dis-
tinct tendency of the posterior portion of the shell to be higher than the anterior,
and thus the shell is not oblicpie. A posterior ridge may be distinct or indistinct.

The hrst species, D. parallelupipedon, (Lea) froms a transition toward the
group of chilensis.

The metropolis of these forms is in the system of the Rio de la Plata, but they
are also found in the coastal streams of southern Brazil (Rio Grande do Sul).

13. DiPL()Dt)N PARALLELOPIPEDON (Lea) (1834).

U7iio parcUlelipipeclon JI’Orbigny, 1843, p. (iOO; Corsi, 1900, p. 447, fig. 30.
Diplodon pamllelipipcdon Simpson, 1914, p. 1275.

Unio pamllelopipedon Von AIartens, 1894, p. 164; Pilsbry & Rush, 1896, p. 30.
Diplodon acutiroslris (Lea) Simpson, 1914, p. 1276.

Diplodo7i pnrallelipipedon and acutirostris Haas, 1913, pp. 22, 23, 52, 53.

Type-locality. — Rio Parana, Province of Corrientes, Argentina.

Other Loealities. — Arroyo del Rosario, Uruguay (to La Plata) (D’Orbigny);
Arroyo de las Vaccas, Uruguay (Gorsi) ; Rio de la Plata, C'olonia, Uruguay (Pilsbry
& Rush); Rio Uruguay, Salto Oriental, Uruguay (Haas); Swamps of Rio Parana
from Buenos Aires to above CVirrientes, Argentina (D’Orbigny); Rio Paraguay
(Von Ihering, 1893, ]). 119); Paraguay (Von Martens); Rio de San Miguel, Prov.
of Chiipiitos, Bolivia, (D’Orliigny).

The last locality deserves special attention, since it is in the Amazons-drainage;
but it is in the most southern extremity of it, close to the divide toward the Para-
guay. It certainly requires confirmation.

New Localities. — Rio Uruguay (in mud), Uruguayana, Rio Grande do Sul,

ortmann: south American naiades.

505

Brazil (J. D, Haseman coll., February 5. 1909). One specimen, male, with soft

parts. Pond along Rio Negro, Santa Isabel, Uruguay (J. D. Haseman coll.,
February 11. 1909). Four complete shells, and five isolated valves; of two of

these soft parts, males.

Distribution. — Drainage of the Rio de la Plata from its mouth and its tributaries
in Uruguay and southern Brazil up to the Rio Paraguay in Paraguay. Also reported
as crossing the divide, and going into the headwaters of the Amazons in Bolivia.
Not known from the Parana above Uorrientes.

This is a species easily recognized by the elongated-siibtraiiezoidal shape,
with the upper and lower margins nearly parallel, by the anterior position of the
beaks, by the rather swollen shell and distinct (although rounded) posterior ridge.
The sides of the disk are rather flattened, and in this respect this species resembles
the cMlensis-gi'owx), intergrading with it to a degree. j\Iy specimens are somewhat
variable in shape, being longer or shorter. None of them shows the lieak-sculpture,
since the beaks are badly eroded in all, except in the specimen from Uruguayana,
where they are only a little eroded, and consequently a little more elevated. But
here also no beak-sculpture can be seen. It must occupy only a very short space
near the beaks, hardly more than 5 mm. The specimen from Urugua3"ana has the
nacre suffused with red (already mentioned by D’Orbigny).

There is not the slightest question that U. acutirostris Lea. is an old, much
eroded, and somewhat distorted specimen of this species. Haas has already sug-
gested this.

Anatomy. — 1 have onl}^ the soft jiarts of male specimens, Imt Lea has already
described them in the case of his D. acutirostris, and has furnished at least some
information about the marsiipium.

Judging from my material, the anal opening is closed above without forming
a supra-anal. Closed part over five times as long as the open, the latter slit-like,
short, shorter than the branchial oiiening. Inner edge of anal indistinctl}" crenu-
lated. Anal and branchial separated by a solid bridge, running a certain distance
inward. Branchial with fine papillae, aliout three times as long as anal. Mantle-
edges not united in front of it. Palpi subtriangular, lower margins convex, pos-
terior margins connected for about one-fourth or one-third of their length.

Gills long and rather narrow. In their posterior part they are of equal width,
or the outer one is slightly wider; anteriorl}^ the inner is much wider. The outer
is considerabl}" narrowed anteriorly, its anterior end being situated near the highest
point of the line of the attachment of the mantle. The inner gill has a straight
margin in the middle, and anteriorly it is onl}^ slightly narrower, its anterior end

506

MEMOIRS OF THE CARNEGIE MUSEUM.

licing immediately behind the palpi. Inner lamina of inner gills entirely con-
nected with abdominal sac. Structure of the gills in the male as usual, but the
interlaminar tissue is unusually well-developed, forming a rather thick layer
chiefly on the inside of the primary limb of the gill; and it has, as usual, short,
interrupted interlaminar connections, elongated in the direction of the gill-filaments,
which in the middle of the gill are few and far apart, while the}^ are a little more
frequent near the ends.

Lea describes the marsupium (of acutirostris) as occupying nearly the whole
length of the inner gill, but no information is given as regards the finer structure.

Color of foot brown or blackish in the distal part, otherwise the soft parts are
whitish.

14. Diplodon charruanus (D’Orbigny) (1835).

Glochidium: Text-fig. 4(/, p. 469.

Unio charruana D’Orbigny, 1843, p. 606, PL 71, figs. 8-11; Pilsbry & Rush, 1896,
p. 81; CoRsi, 1900, p. 447, fig. 31.

Unio faba (as form of charruana) D’Orbigny, 1843, p. 606 (text), as rhuacoica,
PI. 71, figs. 12-14 {in tabula per erroreni, see Explanation of plates, p. 704).

Unio rhuacoica D’Orbigny, 1843, p. 606, PI. 69, figs. 4, 5; Corsi, 1901, p. 450,
hg. 33.

Unio aihiops Lea, Obs., X, 1863, PI. 41, fig. 285 (juv.).

Unio parcus Lea, Obs., Xll, 1869, PI. 33, fig. 77 {juv.).

Diplodon rhuacoicus, charruanus, cElhiops, parcus, Simpson, 1914, pp. 1242, 1243,
1247, 1256.

Diplodon hidcdgoi Haas, 1916, pp. 18, 49, PI. 1, fig. 1.

Diplodon parcus Haas, 1916, pp. 16, 49.

Diplodon for tis IMarshall, 1917, p. 382, PL 52, figs. 1-4.

Type-locality. — Small streams from Maldonado and Montevideo to Las Vacas,
Uruguay (‘'Banda Oriental”).

Other Localities. — Lake Potrero, Maldonado, Uruguay (Pilsbry & Rush) ;
Rio Canelon Grande, Montevideo (D’Orbigny, rhuacoica); Dep. Canelones, Uru-
guay (C’orsi); Rio Miguelcte, Uruguay (Haas, hidcdgoi); Rio Negro, Tacuarembo,
Uruguay; correctly S. Fructuosa, on Rio Tacuarembo, tributary to Rio Negro
(Marshall, /(/rO's) ; Uruguay River (Lea, cetkiops).

New Loccdity. — Pond along Rio Negro, Santa Isabel, Uruguay (J. D. Haseman
coll., February 11. 1909). About twenty-five specimens, seventeen with soft

parts, including males and gravid females.

ortmann: south American naiades.

507

Distribution. — Small streams of the ‘‘Banda Oriental” in Uruguay, from
Maldonado westward, and also in the Rio Negro and the Rio Uruguay.^ ^

An extremely variable form, of which I possess a good set from one locality,
undoubtedly representing young and adult specimens of the same species, so that
I am able to give a rather full account of it. It is very evident that different in-
dividual phases have been previously described as separate species.

Characters of Shell. — Shell of medium size (maximum length according to
D’Orbigny, 70 mm.; my largest is 62 mm.), solid and rather heavy. Outline sub-
trapezoidal, more or less elongated, straight (not oblique), but very variable. The
upper margin may be rather straight (chiefly in young ones), or more or less curved
(in older ones), with or without a distinct posterior upper angle. The posterior
end of the shell is more or less pointed; the position of the point is variable, but
generally rather low, and little elevated above the base-line. The lower margin
is gently curved, often nearly straight in part (chiefly so in old shells), and never
curved up suddenly in its posterior part, but only gently and gradually so, if at
all. The posterior end of the shell is thus distinctly more tapering than the rounded
anterior end, the posterior point lying rather low. The proportion of height to
length of the shell is very variable, ranging from 48 to 65 pr. ct.

Valves quite convex and swollen, hardly flattened upon the sides, but more
convex anteriorly and over the posterior ridge, which is blunt, but more distinct
towards the beaks (and in young shells). Diameter 34 to 50 pr. ct. of length.
Beaks a little inflated, but not very prominent, located at 21 to 29 pr. ct. of length.
Beak-sculpture seen only in my youngest specimens, extending not more than 8
mm. from the point of the beak, consisting of about thirteen radial bars, of which
only the lower ends are seen. They increase little in length posteriorly, are rather
sharp in front, but the longest are somewhat obtuse, while the two last, right upon
the posterior ridge, are again sharp and shorter. In some of my specimens, chiefly
the younger ones, there are some short, oblique wrinkles (one to eight) upon the
posterior slope; in others they are entirely absent. Lunula absent or iiresent,
narrow.

Epidermis with numerous, irregular, finer and coarser, concentric lines; the
finer ones sublamellar on iiosterior slope and towards the margin. Radial sculp-
ture obscure, but present in the shape of fine lines, sometimes more distinct on
the anterior portion of the shell. Color of epidermis dark olive-green to black.

Von Iheriiig (1893, p. 102) reports aihiops from the Ciuahyba drainage in Rio Graiule do Sul,
and a variety (pvracicabana) from the Ui)per Rarand-drainage in Sao Paulo, but these records should be
doubted.

508

MEMOIRS OF THE CARNEGIE MUSEUM.

Old specimens are generally uniformly black; younger ones are dark greenish
olive or brownish olive, sometimes with more or less distinct concentric lighter
(brownish to yellowish) bands.

Hinge-line gently or more strongly convex. Ligamental sinus over the pos-
terior part of the lateral teeth; in larger specimens over the last fifth. Laterals
rather straight in young, curved in older shells, one in right, two in left valve,
their edges somewhat rough. Pseudocardinals normally two in right, and one in
left valve, but often there is a second (posterior) one in the left valve. These
teeth are extremely variable. In younger shells they are compressed and lamellar,
directed obliquely forwards; the posterior tooth of the right valve is always thicker
and higher than the anterior, and they are, chiefly the former, crenulated or denti-
culated. In older shells, the posterior tooth of the right, and the (anterior) tooth
of the left valve may become thicker, more sturniiy, and may be much divided.
The posterior tooth of the left valve, if present at all, may be larger or smaller,
compressed or stumpy.

Cavity of shell and lieaks moderate. Nacre in all my specimens white, iri-
descent posteriorly. Anterior adductor-scar distinct and impressed; anterior
retractor-scar separated from it, small and deep; anterior protractor-scar connected
with it. Posterior adductor-scar less impressed, the posterior retractor-scar forming
an upper process of it. Pallial line distinct. Dorsal muscle-scars a few, situated
in the beak-cavity.

Measurements.

No. 1 Sex.

Length.

Height.

Diameter.

Beaks.

14

34 mill.

20.5 mm. =60 pr. ct. of L.

12

mm. =35 pr. cl. of L.

at 10

mm. =29 pr. ct. of L.

12

48

26.5 “ =55

20

“ =41

10

“ =21

c

cf

54

35 “ =65

27

“ =50

14

“ =26

9

9

54

30.5 “ =56

24

“ =44

13

“ =24

Half shell

58

31 “ =53

26

“ =45

13

“ =22

3

9

61

35 “ =57

26

“ =42

17.5

“ =29

z

62

35 “ =56

27

“ =44

15

“ =24

For comparison I give here previous measurements.

Length.

Height.

Diameter.

Beaks.

charruanus, D’Or-
bigny’s tc.xt:
rhuacoicus, D’Or-

70 mm.

61 pr. ct. of L.

45 pr. ct. of L.

bigny’s text;
faba\ {rhuacoicus),

63 mm.

51 pr. ct. of L.

43 pr. ct. of L.

D’Orbigiiy’s fig.

43

20.5 mm. =48 “

15 mm. =35 “

12

mm. =28 pr. ct. of L.

cclhiops. Lea’s fig.

.53

28 “ =53

18 “ =.34

12

“ =23

parcus, Lea’s fig.

IS “ =50

13 “ =36

10

“ =28

hidalgoi, Haas:

63

39 “ =62

27 “ =43

66.5 “

42 “ =63

27 “ =41

fortis, Marshall;

66

37 “ =56

27 “ =41

ortmann: south American naiades.

509

Remarks. — Among our specimens the single valve comes nearest to the measure-
ments given for U. rhuacoicus. D’Orbigny himself admits that this may be only
a more elongated form of charruanus, and his faba, united by him with charruanus,
is even more elongated. There is no reason for keeping rhuacoicus separate, since
the proportional length is said to be the only difference.

The variations in the shape of the shell are, indeed, very great. Younger
specimens are, however, more uniform; they are more elongated, and their normal
shape is well rendered in D’Orbignj^’s figures of faba (PI. 71, figs. 12-14), and in
the figures of octhiops and parcus. The latter undoubtedly is a young shell, but
(Ethiops also belongs here. Specimens like those measured under Nos. 12 and 9
resemble this very much, except that they are a little more swollen, and our smallest
(No. 14) looks very much like parcus, but it is less elongated.

In the description and the figures of D’Orbigny only the color of the epidermis
is not exactly as in our specimens. D’Orbigny describes it in charruanus as well as
rhuacoicus as brownish green, and figures it as lighter or darker lirown, while the
color oi faba (PI. 71, figs. 13, 14) is blackish, agreeing better with our specimens.
However, our young specimens have a color, which may be called brownish green,
or rather “olive-brown.” Lea’s octhiops is said to be black.

The chief characters of this species thus seem to be the subtrapezoidal, rather
elongate, straight shell, with a moderately sharp posterior point, located only
little above the base-line; the somewhat swollen shell, with a moderate and blunt
posterior ridge, and the lirown to blackish color of the epidermis. The chief varia-
tions are found in the length of the shell, and the somewhat higher or lower position
of the posterior end.

There is no question that hidalgoi Haas and fortis Marshall are this species.
The former is founded upon two specimens, the latter upon a single one, which
certainly represent individual phases. Specimens greatly resembling these are
among my material.

Anatomy. — I have the soft parts of eleven males and six gravid females, three
with eggs, three with glochidia, one of these discharging.

Anal opening slit-like, closed above, the closed part about four times as long
as the opening; the latter shorter than the branchial opening. The two openings
are separated by a solid mantle-connection. Branchial with small papillae. Palpi
subtriangular, their posterior margins connected near the base. Gills posteriorly
of about the same width, but anteriorly the inner is wider, and its anterior end is
immediately behind the palpi. Inner lamina of inner gill entirely connected with
abdominal sac. Non-marsupial gills with scattered, short, interlaminar connec-

510

MEMOIRS OF THE CARNEGIE MUSEUM.

tions. The marsupial part of the female occupies a large portion of the inner gill,
leaving about one-fourth of the gill free in front, and a smaller part free behind,
thus gravitating slightly toward the posterior part of the gill. The interlaminar
connections could not be distinctly observed by me on a lateral view, since no barren
females are at hand; l)ut from sections it was possible to infer that they form in-
terrupted sejita forming a system of intercommunicating water-tubes. The eggs
and glochidia hll the water-tubes and the perforations of the septa in a mass, which
is not conglutinated and divided into placenta;.

The glochidium (Text-fig. 4(/, p. 469) has the characteristic triangular shape,
somewhat oblicpie, with the point situated below the posterior end of the upper
margin (like the figure of the glochidium of U. peculiaris, See Lea, Obs. XII, 1869,
PI. 34, fig. 80). This point does not possess a hook. Size of glochidium: L. 0.31,
D. 0.26 mm.

I have examined the glochidia of three specimens; one of these had the mar-
supium largely empty, and thus it appears to have been discharging. Yet no hooks
were seen. But it may be that the discharge in this case was premature, that
none of the glochidia were mature, and that the hooks might have developed later.
This can be decided only by investigating more material.

15. Diplodon piceus (Lea) (1860).

Anatomy of gills: PI. XLVI, fig. 2; glochidium: Text-fig. 4/i, p. 469.

Unio piceus Lea, Obs., X, 1863, PL 41, fig. 287.

Diplodon piceus Simpson, 1914, p. 1244; Haas, 1916, pp. 15, 49.

Type-locality. — Uruguay River.

Other Localities. — Rio Uruguay, Salto Oriental, Uruguay (Haas); Rio Migue-
lete, Uruguay (Haas).

Localities Represented in the Carnegie Museum. — Rio Uruguay (in mud),
Uruguayana, Rio Grande do Sul, Brazil (J. D. Hasernan coll., February 5, 1909)
eight specimens, seven of them with soft parts, males and gravid females. Arroyo
Miguelete, Montevideo, Uruguay (J. D. Hasernan coll., February 17,1909) one
specimen.

Distribution. — Positively known from the Uruguay River, and from a small
coastal stream (R. Miguclete) near Montevideo, and probably more widely dis-
tributed in the “Banda Oriental” in Uruguay. It possibly may be only a form of
charruanus. Corsi does not mention it from Uruguay.

A species closely allied to D. charruanus, and very near to it in its dimensions,
except that it does not show the same extremes of variation. It is, however.

ortmann: south American naiades.

511

shorter on the average (H. 59 to 63 pr. ct. of L., while D. charruanus varies from 48
to 65 pr. ct.). In addition D. piceus is not so subtrapezoidal in outline, but rather
subovate, which is brought about by a stronger curve of the lower margin, which
ascends much more distinctly posteriorly, so that the posterior end of the shell is
more elevated above the base-line. At the same time the posterior end is rather
blunt and rounded, and not so pointed as in D. charruanus.

All other characters of the shell are similar to D. charruanus. The radial
sculpture of the epidermis is more distinct. The inside of the shell is also similar,
but the pseudocardinals are simpler, always elongate and compressed, and there is
always only one in the left valve. They are not much cut up, but only crenulated
and rugose. Nacre white, though one of my specimens has purplish blue in the
cavity, probably a discoloration.

Beak-sculpture, as far as can be seen in smaller shells, similar to that of D.
charruanus. But there is a very small specimen, which is only 10 mm. long, among
them, which is doubtfully referred here. In this case the sculpture consists of
fifteen radial bars, of which the eighth and ninth unite at their lower ends. The
anterior bars are shorter, but there is not much difference in this respect from the
posterior bars. The bars are rather sharp and fine, but those upon the umbonal
ridge are slighth^ thicker, and the two last, upon the posterior slope are very fine
and shorter.

Measurements.

No.

Sex.

Length.

Height.

Dmmeter.

Beaks.

Uruguayan

a.

1

9

.30 mm.

19 mm

= 63 pr. ct. of L.

11.5

mm. =38 pr. ct. of L.

at 8.5

mm. =28 pr. ct. of L.

91

9

35

20.5 ‘‘

= 59

13

“ =37

10

“ =29

5

d'

51

31

= 61

21

“ =41

15

“ =29

4

9

51.5 “

30

= 58 “

20.5

“ =40

15

“ =29

7

9

64

38

= 59

28

“ =44

18

“ =28

Montevideo.

57 “ ■

35.5 “

= 02

25.5

“ =45

16

“ =28

Lea’s fig.

47

28

= 60

19

“ =40

12

“ =26

Remarks. — There is not the slightest doubt that my siiecimens represent the
U. piceus of Lea, but the question of its possible identity with D. charruanus must
be left undecided, also that of other jiossible synonyms {lepiclus Lea, and firmus
Lea). I cannot unite these for the. present on account of certain jieculiar structures
in the anatomy, which will be pointed out presently.

Anatomy. — I have the soft parts of one male, one barren female, and five
gravid females, three of the latter with eggs, and two with glochidia, but in one
only are the glochidia mature.

The structure is entirely like that of D. charruanus. With regard to the mai'-

512

MEMOIRS OF THE CARNEGIE MUSEUM.

supium (PI. XLVI, fig. 2) it is to l:)e remarked that it occupies in larger specimens
a somewhat greater ])ortion of the inner gill, leaving anteriorly as well as posteriorly
less than one-foiirth of the gill free. In smaller females the marsupial part is
relatively smaller. The interlaminar connections form distinct, but interrupted,
septa. The connections stand very close, and the septiform structure prevails
throughout the marsupium, and very little is seen of a transverse or reticulate
arrangement. In some parts, chiefly towards the margin, the septa become more
or less continuous. Thus there are here rather distinct, but intercommunicating
water-tubes (ovisacs), which are filled, when charged, with masses of eggs connected
with each other through the communications, so that no i)lacenta-like arrangement
is observed.

Glochidium (Text-fig. 4/;, ]i. 469). — Higher and more upright than that of D.
charriianus’, L. and H. about the same, 0.28 to 0.29 mm. Thus it is more like that
of D. Jirmus figured by Lea (Obs., XII, 1869, PI. 34, fig. 82), but not quite as iqiright.
At the ])oint of the lower margin there is a hook of the usual shape, about 0.09 mm.
long. Put such hooks are present in only one of my specimens, in the other the
glochidia are too immature.

It would be quite remarkable if in two species, so closely allied as D. charruanus
and piceus, the glochidia should tliffer so fundamentally, that in one there are
hooks, and in the other not. Put judging from my material, this is the case. How-
ever, it must be enq^hasized again that my material is scanty, and possibly in the
case of D. charruanus I do not at all have rijie glochidia.

16. Diplodon uruguayensis (Lea) (1860).

Unto Uruguay ensis Lea, Obs., X, 1863, PI. 45, fig. 298.

Diplodon uruguayensis Simpson, 1914, p. 1234.

Unto apprimus Lea (1866), Obs., XH, 1869, PI. 33, fig. 78.

Type-locality . — Uruguay River.

New Locedity. — Pond along Rio Negro, Santa Isabel, Uruguay (J. D. Haseman
coll., Feliruary 11, 1909). Five complete specimens, and several odd valves,
two specimens (male and female) with soft parts.

Distribution. — Known only from Rio Uruguay and Rio Negro.

The synonymy and affinity of this species is obscure. Simpson thinks that it
is close to D. ivyrnani, but the latter is a compressed shell, while uruguayensis is
much swollen. U. appriynus also is a swollen shell. Lea has already suggested
that this is closer to uruguayensis, but that it differs chiefly in the hinge-teeth.

ortmann: south American naiades.

513

which are more lamellar and compressed in the latter (a smaller shell), and more
stumpy and cut up in apprimus. My material shows that the character of the
hinge-teeth changes with age. In general my specimens correspond in size and
shape to Uruguay ensis, but the larger ones have more stumpy teeth, and thus I
believe that apprimus is an old specimen of uruguayensis. Both Simpson and
Haas (1916, p. 12, 47) unite apprimus with wymani, which cannot be correct on
account of the difference in obesity. I think wymani belongs to lacteolus (See
below) .

However, it is quite possible that all these forms are variations of one and
the same species, and then, of course, our general arrangement must be changed.
As regards the present form I can only say that it looks like a very large and heavy
charruanus, the shell being rather elongated, subtrapezoidal, and much swollen.
In lacteolus, the shell is higher and more ovate, and mucli more coinjiressed. In
none of my specimens is the beak-sculpture seen.

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

a

73 mm.

50

mm. =68 pr. ct,. of L.

31 mm

= 43 pr. ct. of L.

at 19 mm

= 20 pr. ot. of L.

1

73.5 “

47

“ =64

37 “

= 50

20 “

= 27

b

78

52

“ =67

40 “

= 51

21 “

=27

2

9

78

52

“ =67

33 “

= 42

19 “

= 24

c

79

47.5

“ =00

32 “

=41

20 “

= 25

uruguayensis

70

45

“ =64

30 “

= 43

apprimus'^

101

06

“ =65

40 “

= 40

Anatomy. — Soft parts of a male and a barren female at hand.

Color of soft parts : distal part of foot dark gray, this color sharply marked off
from the whitish basal part.

Anal closed above; closed part about four times as long as the open part,
which is slit-like, and shorter than the branchial, and separated from it by a solid
mantle-connection. Branchial opening with small, but distinct, papillae. Palpi
rather large, subtriangular, with lower margins strongly convex; posterior margins
connected at base.

Gills long and moderately wide, the inner one much wider than the outer
anteriorly, its anterior end immediately behind the palpi. Inner lamina of inner
gill entirely connected with abdominal sac. Non-marsupial gills with scattered,
short, interlaminar connections. In the female a large section of the inner gill
is marsupial, about one-fourth of the gill remaining non-marsupial at the anterior

The measurements of uruguayensis and apprimus are tliose given by Simpson for the respective
types. Those for apprimus are given by him under wymani (p. 1231).

514

MEMOIRS OF THE CARNEGIE MUSEUM.

end, and less than that at the posterior end. The interlaminar connections of the
marsupium are developed as interruiited septa, forming communicating water-
tubes. The connections are short, and are also arranged in irregular cross-rows^
but towards the edge of the gill, the sei)tiform arrangement is quite distinct.

17. Diplodon hildvE Ortmann, sp. nov.

Shell: PI. XXXVI, fig. 7; PI. XXXVII, figs. 1, 2, 3; Anatomy of gills: PI. XLVI,

fig. 3; Glochidium: Text-fig. 4f, p. 4G9.

Type-locality. — Rio Jacuhy, Cachoeira, Rio Grande do Sul, Brazil (J. D.
Haseman coll., January 26, 1909). Type-set: Carnegie Vluseum, No. 61.5864.
Fourteen s]iecimens, males, barren and gravid females, soft parts of all in alcohol.

There were six additional specimens in the original lot.

Descriptio7i of the Shell. — Shell rather small (maximum length 46 mm.), rather
solid. Outline subellijitical or indistinctly subtrapezoidal, height 57 to 63 pr. ct.
of length. Upiier margin straight or gently convex, forming a blunt angle with
the iiosterior margin. Posterior margin obliquely descending, straight, or gently
convex, forming a blunt point with the lower margin ; this ])oint situated distinctly
above the base-line. Lower margin rather regularly convex, ascending anteriorl}^
and iiosteriorly, its lowest jioint situated between beaks and posterior end of
ligament (rather median). Anteriorl}' the margin is regularly rounded. Anterior
end of shell not much narrower than the iiosterior, which tapers to the blunt pos-
terior point, so that the shell is rather regularly elliptical, with the posterior end
subiiointed, the angle of the posterior uiiiier margin giving a suggestion of the
subtrapezoidal shape.

Valves rather regularly convex, greatest diameter slightly behind the middle,
but distinctly in front of the posterior ridge, which is very blunt and broad. Pos-
terior slope somewhat compressed, sometimes with a trace of a radial furrow.
Sides of disk less convex, but not at all flattened. Diameter 34 to 44 pr. ct. of
length, so that the shell appears as moderately swollen. Beaks a little inflated,
but not very ]irominent above the hinge-line, located at 23 to 28 pr. ct. of the
length. Beak-sculpture seen only in the smallest specimens, and only the lower
part of it. There are aliout fourteen radial bars, the anterior rather sharp, the
posterior located just in front of the posterior ridge, broader (at least their lower
ends) and the latter are a little longer than the former. The last two or three
bars are again finer and shorter. The longest bars are about 8 mm. long. There
may be fine, short, oblique wrinkles upon the posterior slope, but these are distinct
only in a few siiecimens. Lunula short, very narrow, or practically absent.

ortmann: south American naiades.

515

Epidermis shining, chiefly so in the middle of the disk and in younger speci-
mens, but with unequal concentric wrinkles and striae, which become somewhat
sublamellar on the posterior slope and near the margins. Radial sculpture jiresent,
consisting of irregular, often interrupted lines, chiefly upon the anterior part of
the shell, often appearing as ‘‘scalariform stripes’’ (radial rows of fine, short, con-
centric wrinkles). Color of epidermis light to dark brown; in young specimens it
is a beautiful, shining, golden brown, lighter in the middle of the shell and towards
the beaks (and sometimes here with light greenish shades). In older shells the
color is chestnut-brown to dark brown. In some specimens there are indications
of darker concentric bands. No traces of color-rays, except one or two very faint
dark rays upon the posterior slope (seen only when held up against a strong light) .

Hinge-line very gently curved. Ligamental sinus over the posterior fourth or
third of the lateral teeth (more posterior in old shells, which shows that the ligament
is comparatively longer in them). Lateral teeth gently curved, long, one in right,
two in left valve, finely rugose. Pseudocardinals obliquely directed forward and
downward, normally two in right, one in left valve, comjiressed, but not very
long, their edges crenulated and serrated. The posterior in the right valve more
elevated than the anterior, and often thicker. Sometimes there is a trace of a
second posterior tooth in the left valve, but this is always small.

Cavity of shell and beaks moderately deep. Nacre whitish, shining, in the
largest specimens the thickest parts (toward the front of the shell) have a faint
rosy hue. Muscle-scars of the typical shape; the anterior adductor-scar is rather
deep, the retractor-scar separated from it, round and deep, the protractor-scar
connected with it; the posterior adductor- and retractor-scars are united. Pallial
line distinct. An irregular, longitudinal row of dorsal scars in the beak-cavity.

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Figured.

d...

d'

25 mm.

14.5 mm

= 58 pr. ct. of L.

8.5

mm. =34 pr. et. of L.

at 7

mm. =28 pr.

ct. of L.

PI. XXXVII, 6g. 1.

c. . .

9

32.5 “

19.5 “

= 60

11.5

“ =35

7.5

“ =23

' k. . .

d

37.5 “

23.5 “

= 63

15.5

“ =41

10

“ =27

p.. .

9

40

24

= 60

16.5

“ =44

10.5

“ =26

q.. .

d

43

24.5 “

= 57

17

“ =40

10

“ =23

PI. XXXVI, fig. 7.

15.. .

9

46.5 “

27

=58

19

“ =41

11.5

“ =25

PI. XXXVII, fig. 2.

Remarks. — I have been unable to recognize this species in any of the published
descriptions. It is a rather beautiful shell; marked by its small size, nearly
regularly elliptical outline, with no obliquity, rather swollen valves, and with a
peculiar, shining, golden brown color when young. It resembles, to a degree,
D. charruanus, but differs from it chief!}" in size, more regular shape, and the gloss
of the epidermis.

516

MEMOIRS OF THE CARNEGIE MUSEUM.

Very few species of Diplodon are known in Rio Grande do Sul, from the drainage
of the CJiiahyba (to which the Rio Jacuhy belongs) and all have been only incident-
ally mentioned by Von Ihering (1893). Under U. wthiops piracicabana (L c., p.
102) he reports that U. cethiops is found in the Guahyba River. I have been unable
to discover this species, which is identical with charruanus, in the material col-
lected by Haseman in this system, and I am sure that the present form is not the
one which Von Ihering calls aihiops. From his casual remarks it is seen that the
latter has a shallow depression on the disk (/. c., p. 104), and nothing of the kind
is seen in D. hilda\

Anatomy. — I have two barren females, and three gravid females with glochidia.
Of the latter, one had only very few glochidia in the marsupium, but some were
in the suprabranchial canals, and thus it was evidently discharging. The rest
of my specimens are males.

Color of soft parts whitish; distal part of foot grayish black.

Anal 0}iening closed above; closed part about four times as long* as the open
part, the latter slit-like, slightly shorter than the branchial; the latter has small,
but distinct ])apilhe. Anal and branchial oiienings separated by a solid mantle-
connection. Palpi moderate, sulitriangular, posterior margins connected at base.

Gills long and rather wide. They are about eiiually wide posteriorly, but the
outer is sulitriangular and narrows anteriorly, while the inner does not, and re-
mains as wide as ]iosteriorly, with its anterior end immediately behind the palpi.
Inner lamina of inner gill entii-ely connected with abdominal sac. The non-
marsupial gills with scattered interlaminar connections. In the female, the mar-
S'upium (Plate XLVI, fig. 3) is located in the inner gill, but anteriorly about one-
third of the gill remains non-marsupial, posteriorly much less, about one-fifth of
it or less, so that the marsupial iiart is located distinctly more backward in the
gill, occupying about half of its length. When charged, the marsupium forms a
slightly swollen patch. In young specimens the marsupium is smaller. Inter-
laminar connections in the marsupium forming septiform rows, with a tendency
to fall also into irregular transverse rows. The transverse arrangement prevails
near the base and in the middle of the gill, here and there with a suggestion of a
([uincuncial disposition; the septiform arrangement is found toward the margin
of the gill.

Glochidium (Text-fig. 4t, p. 469) subtriangular, longer than high, with a ventral
point, slightly oblique, the point being vertically under the posterior end of the
upper margin. There are no hooks. In one of my sjiecimens, the glochidia are
not margined (not mature); in the two others they are margmed, with a narrow

ortmann: south American naiades.

517

rim around the anterior-lower-iiosterior margin, representing, apparently, the first
rudiments of the postembryonal shell. Size (without rim): L. 0.29 to 0.30; H.
0.26 mm.; (with rim): L. 0.34 to 0.35; PI. 0.28 to 0.29 mm.

The hookless glochidium, provided with a rim or margin, is highly interesting
in view of the fact, that this structure has not been observed in other species of the
charruanus-group .

5. Group of Diplodon lacteolus.

Like the fourth group (that of charruanus), but shell higher and shorter,
subtrapezoidal to ovate, chiefly so when j^oung, conijiressed or somewhat swollen.
Beak-sculpture fine or a little heavier and better develoi)ed, but not covering a
large part of the shell.

This group stands close to that of D. charruanus. The greater height of the
shell is chiefly evident in the young shell, which may be slightly oblique. But
the older shells are also shorter and higher than in the sjiecies of the charnianus-
group, although they generally are less elevated than younger shells. They are
not oblique, and have a rather regular, bruadly ovate, or subelliptical outline.

18. Diplodon burroughianus (Lea) (1834).

Anatomy of gills: Plate XLVI, fig. 4.

Unio burroughianus Lea, Obs., I, 1834, PI. 10, fig. 27; D’Orbigny, 1843, p. 609;

Von Martens, 1894, p. 164; Corsi, 1901, p. 450.

Diplodon burroughianus Simpson, 1914, j). 1271.

Type-locality. — Rio Parana, Province of Corrientes, Argentina.

Other localities. — Small rivers of the Banda Oriental in Uruguay (D’Orbigny) ;
Montevideo (D’Orbigny) ; swamjis along the Parana from Buenos Aires to above
Corrientes (D’Orbigny); Paraguay (Von Martens).

It also occurs near Santa C’ruz de la Sierra in Bolivia (D’Orbigny) which is
in the drainage of the Amazons.

New Locality. — Pond near the Rio Negro, Santa Isabel, Uruguay (J. D. Hase-
man coll., PVbruary 11, 1909). Three complete specimens, two of them, male
and female, with soft parts, and four odd valves.

Distribution. — Drainage of Rio de la Plata, including the small streams of the
Banda Oriental, Rio Negro, and the Parana and Paraguay to Paraguay, and possibly
also in the Amazon-drainage in Bolivia. The latter part of the range, however,
should be confirmed.

Some of my specimens, chiefly the female with soft parts, agree very well with

518

MEMOIRS OF THE CARNEGIE MUSEUM.

hurroughianus in shape, dimensions, and color. Others differ more or less, chiefly
in having the posterior point of the shell less elevated above the base-line. Thus
they approach other species described by Lea as jiiger (1860) and ampullaceus
(1869), both from the Uruguay River (See also ampullaceus, Haas, 1916, pp. 11,
47). My material is not sufficient to work out the synonymy.

Measurements.

Sex.

Lengtli.

Height.

Diameter.

Beaks.

9

54 mm.

38 mm.

= 70 pr.

?.t. of L.

23.5

mm. = 44 pr. ct. of L.

at 13 mm. = 24 pr. ct. of L.

cf

56.5 “

39 “

= 73

i (

27.5

<< _49 “

15 “ =27 “

60

44 “

= 73

( (

26

“ =43

16 “ =27 “ (Lea’s figure)

100

62

1 (

44

(D’Orbigny)

94

64 “

= 68

((

40

“ =43 “

(Von Martens)

88

58 “

= 66

( (

35

“ =40

(Simpson)

71

43 “

= 61

“

26

“ =37

Do. y

The two specimens measured liy Simpson do not include the type. They are
more compressed than any others.

Anatomy. — The soft parts of a male and a female are at hand.

Color of soft parts whitish, distal part of foot grayish.

Anal opening slit-like, closed above, short, shorter than the branchial opening,
separated from the latter by a connection of the mantle-margins. Branchial
opening short, its inner edge with small, but distinct paiiillse. Palpi subtriangular,
moderately large, lower margins convex, posterior margins united at base for a
short distance.

Gills of the usual shape; the inner the wider, chiefly in front, its anterior end
close behind the palpi. Structure of non-marsupial gills as usual. Marsupial
portion in the female (PI. XLVI, fig. 4) located in the inner gill, leaving about one-
third at the anterior end, and about one-fifth at the posterior end free, so that the
marsuiiium distinctly gravitates toward the posterior part of the gill. Interlaminar
connections arranged in intermitted septa, and irregular, transverse rows, here
and there quincuncial. This irregularly reticulate structure prevails throughout
the marsupium, and the septiform arrangement is obscure.

19. Diplodon lacteolus (Lea) (1834).

See; Diplodo7i lacteolus (Lea) and D. wymani (Lea) (1860) Simpson, 1914, p. 1226,
1230.1®

Lea liimself (1834, p. 90) lias identified the type of Lamarck’s U. delodonta (1819) with his lacteolus
(1834, p. 40), but not until after the latter name had been published in a satisfactory way, while the
original description of delodonta is absolutely unidentifiable. Thus, according to the rules, lacteolus
must be used, as Simpson has done.

ortmann: south American naiades.

519

The following references should be added to those given by Simpson.

Unio delodonta D’Orbigny, 1843, p. 605; Corsi, 1901, p. 449.

Diplodon wymani Haas, 1916, pp. 12, 47.

As to the synonymy compare Simpson. His D. wiymayii only in part belongs
here. D. apprimus Lea, united bj^ him with wymani, is different (more swollen,
see above p. 512). On the other hand, I cannot distinguish the real wymani from
lacteolus, and Von Ihering (1893, p. 117) also unites them.

Type-locality. — Rio de la Plata.

Other Localities. — Stream, Villa del Cerro, Montevideo, Uruguay (D’Orbigny);
Rio Uruguay (Lea, wymani) ; Rio Uruguay, Las dos Hermanas Islands, Uruguay
(D’Orbigny); Rio Uruguay, Salto Oriental, Uruguay (Haas); Rio San Salvador
(trib. to Uruguay), Soriano, Uruguay (Corsi); Buenos Aires (D’Orbigny); Rio
Batel and Rio Corrientes, Province of Corrientes, Argentina (D’Orbigny).

New Localities. — Pond near Rio Negro, Santa Isabel, Uruguay (J. D. Haseman
colL, February 11, 1909). One specimen, young. Brooklet, two miles north
of the City of La Plata, Argentina (Dr. W. J. Holland coll., September 24, 1912).
Two specimens.

Distribution. — Lower La Plata and Parana systems, upward to the province of
Corrientes; also in the lower Uruguay and Rio Negro, and small tributaries of the
La Plata in Argentina and Uruguay.

My specimens from La Plata are typical. The young specimen from Santa
Isabel is interesting in having a rather high shell, which is slightly oblique, but
comparison with the other specimens shows that, according to the growth-rests,
the latter had the same shape when young. This young shell also exhibits the
comparatively heavy beak-sculpture, with the radial bars running down the shell
about 13 to 15 mm. Traces of these are also seen in my larger specimens.

Measurements.

Length.

Height.

Diameter.

Beaks.

Santa Isabel

50

nim.

36.5 mm

=73 pr. ct. of L.

18 mm. =36 pr. ct. of L.

at 13 mm

= 26 pr. ct. of L.

La Plata

85

61

= 72 ■

32 “ =38

^1 “

= 25

Do

87

63

= 72

34 “ =39

22 “

= 25

lacteolus (Lea’s figure)

80

51

= 64

31 “ =39

23 “

= 29

Do. (Ace. to Simpson)

80

50

= 63

30 “ =38

Do. Do.

72

42

=58

26 “ =36

Do. Do.

83

56

= 67

35 “ =42

delodontus (D’Orbigny)

85

70

38

wymani (Lea’s figure)

75

49 “

= 65 -

28 “ =37

19 “

= 25 . “

The first measurements given by Simpson apparently refer to the type of
lacteolus, although they do not fully agree with Lea’s figure. The second measure-

520

MEMOIRS OF THE CARNEGIE MUSEUM.

ments given by him belong to a specimen which is exceptionally low. The measure-
ments given by Simpson for wymani do not belong to this species, but to apprimus
(See above, p. 513, footnote 18').

I have a suspicion that D. felipponei Marshall (1917, p. 381, PI. 50, figs. 1-3,
PI. 51, fig. 1) from Maldonado and other places in Uruguay, is also this species.
The height as given is 70 to 72 pr. ct. of the length, and the general shape and
other iiarticulars agree, as for instance the hinge-teeth. But, according to the
figure, felipponei is less pointed behiinl, and the diameter is a little greater (40 to
46 pr. ct. The color also (yellowish-chestnut) is not exactly like lacteolus, which
has chestnut-olive-green tints. Thus 1 leave this question undecided.

20. Diplodon mogymirim Ortmann, sp. nov.

Shells: PI. XXXVIl, figs. 4, 5, 6, 7; Anatomy of gills: PI. XLVI, fig. 5; Section of
gills: PI. XLVIll, fig. 2; Glochidinm: Text-fig. 4/v, p. 469.

Type-locality. — Creek near Mogy Mirim, Sao Paulo, Brazil, tributary to Rio
Mogy Guassu and Rio Grande, upper Parana-drainage. J. D. Haseman coll.,
August 28, 1908. Type-set: Cam. Mus. Cat. No. 61.9260, fourteen specimens, males,
barren and gravid females, all with soft parts. (The original lot contained over
one hundred specimens.)

Of all the species of Diplodon known from the upper Parana-drainage in Sao
Paulo only one described by Von Ihering resembles this in shape: Unio greejfeanus
(Von Ihering, 1893, p. 96, PI. 4, fig. 8). However, the color of the epidermis of
the latter is descrilied as being dark green to blackish, and this does not at all
fit our specimens, which are brownish black, without any distinct greenish tints.
Moreover, the dimensions given for the two specimens described by Von Ihering)
although falling within the range of variation of my specimens, are rather extreme,
and do not represent the normal condition of our species. The height is 61 and
64 pr. ct. of the length, while in my material this proportion varies from 62 to 74
pr. ct.; and the diameter is 34 and 33 pr. ct., while it ranges, in my specimens, from
32 to 42 pr. ct. U. greejfeanus comes from the Piracicaba River (Campinas and
Piracicaba), while our specimens lielong to the Rio Grande drainage.

From the latter, and especially from Rio Mogy Guassu, at Jaboticabal (not
“Taboticabal” as printed), Simpson (1914, p. 1250) has described Diplodon trivialis.
Dimensions and descriiition agree with our specimens to a degree; but again the
color is different, being described as black or dark brown, and tinted green when
rubbed, while in our specimens, when rubbed (cleaned), the epidermis is of a peculiar

ortmann: south American naiades.

521

golden brown, without green. Also the epidermis is not at all ‘‘cloth like,” as
described in trivialis (when fresh), and the description of the pseudocardinals of
the left valve does not agree at all. There are said to be two pseudocardinals,
the anterior one sometimes feeble, while in our specimens there is generally only
one well-developed, and this is the anterior, and if there is a smaller second pseudo-
cardinal, this is the posterior. The figures of trivialis given by Marshall (1917,
p. 386, PL 54, figs. 5-8) show also that the outline is different, being evenly rounded
behind. Among mj^ numerous specimens there is not a single one which shows this
character, and thus I cannot identify them with D. trivialis.

Description of Shell. — Of moderate size (maximum length 68 mm.), rather
solid. Outline short subelliptical or subovate, or subrhomboidal, when young.
Height from 62 to 74 pr. ct. of length. Upper margin nearly straight when young,
more or less curved when old, in the first case forming an angle with the obliquely
and rather steeply descending posterior margin, in the latter case passing into it
more or less gradually. Posterior margin gently concave, straight, or gently
convex, forming a rounded angle with the lower margin, which may be more dis-
tinct in older shells. This posterior point is more or less elevated above the base-
line. Lower margin gently and regularly curved, in older shells more nearlj^
straight in the middle. Anterior end of shell slightly narrower than the posterior
in young shells; in old shells this may be reversed. Thus the shell is, when young,
more subrhomboidal, with an upper posterior angle (somewhat subalate), and,
when old, the shell becomes subelliptical or subovate, with the posterior end a
little tapering.

Valves rather regularly and evenly convex, sides not distinctly flattened.
Greatest diameter a little anterior to the middle. Posterior ridge present, but
rounded and indistinct, often (chiefly in young specimens) marked by a shallow
radial groove running down the posterior slope, which thus appears as compressed
and slightly elevated toward the upper-})osterior margin (subalate). Diameter
32 to 42 pr. ct. of the length, so that the shell is rather compressed. Beaks not
inflated and not much elevated, located at from 25 to 31 pr. ct. of the length. Beak-
sculpture consisting of fine and short radial bars, hardly more than 5 mm. long,
fifteen to eighteen in number, those in the middle converging at their lower ends
(one or two pairs). They are hardly longer upon the posterior ridge, and not ap-
preciably thicker. A few oblique wrinkles may be present upon the posterior
slope. In most cases the beak-sculpture is entirely oliliterated by erosion of the
beaks, and in general it is fine, short, and poorly developed. Lunula present,
narrow in young specimens, wider in old ones, but very variable.

522

MEMOIRS OF THE CARNEGIE MUSEUM.

Epidermis somewhat shining, with unequal, irregular, concentric wrinkles,
more crowded and sublamellar upon the posterior slope and near the margins,
chief!}" in older shells. - Radial sculiiture may be present, but indistinct, visible
chiefly upon the anterior part of the shell as ‘‘scalariform” stripes. Color of epi-
dermis from yellowish brown to dark brown and blackish, but without any dis-
tinct traces of greenish tints. In young specimens, when well cleaned, the color
is generally yellowish or golden brown in the middle of the disk, shading to chestnut-
lirown toward the ends and margins. Older shells are more uniformly chestnut-
brown to blackish brown (often coated with a dull black-brown deposit). Slight
traces of darker brown concentric bands are rarely present.

Hinge-line generally distinctly curved. Ligamental sinus over the posterior
half or third of the laterals, generally very indistinct in old specimens. Lateral
teeth curved (less so in young shells), moderately long, one in right, two in left
valve, their edges and sides corrugated in old shells. Pseudocardinals directed
obliquely downward and forward, compressed and lamellar, thin in young shells,
thicker in old ones. The right valve has nearly always two of them, equally high,
but the anterior narrower. The left valve has mostly only one; but there may be
a small and short posterior one, and even a small and narrow anterior one. In
one specimen (one out of over one hundred) , the pseudocardinals are exactly re-
versed: one in right, two in left valve, while the laterals are normal. The edges
of the iiseudocardinals are rugose and crenulated, but not dissected.

C’avity of shell and beaks moderate. Nacre whitish, l:)ut very generally parti-
ally discolored, with irregular yellowish, brownish, or grayish spots. Anterior
adductor-scar imjiressed, rounded or subtriangular; anterior retractor-scar separated
from it, rounded or irregularly oval, imjiressed, but not remarkably deep; anterior
protractor-scar connected with adductor-scar. Posterior adductor-scar ovate or
subtriangular, less deeply impressed; posterior retractor-scar generally separated
from it, l)ut sometimes only indistinctly so. Pallial line distinct. Dorsal scars
few and irregular, in beak-cavity, forming an indistinct longitudinal row.

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Eiguretl.

49. . .

&

31 iniii.

22

mill

= 71 pr.

cl. of L.

11

nmi. =35 pr. ct. of L.

at

8.5

mm. =27 pr. ct. of L.

12.. .

d'

39

26.5

= 08

12.5

“ =32

11

“ =28

PI. XXXVII, fig. 6.

18.. .

9

40.5 “

30

=74

15

“ =37

11

“ =27

9.. .

d

53

30

= 08

20

“ =38

13.5

“ =25

PI. XXXVII, fig. 5.

41.. .

9

03

39

= 02

2(i

“ =41

17

“ =27

5.. .

9

04.5 “

40

= 02

27

“ =42

17

“ =20

34.. .

9

08

43

= 03

25.5

“ =38

21

“ =31

16.. .

9

68

45

= 00

20

“ =38

18

“ =26

ortmann: south American naiades.

523

Remarks. — This species somewhat resembles D. lacteolus, but is considerably
smaller, has finer and shorter beak-sculpture, and, when old, has more simple,
less dissected pseudocardinals, of which those of the right -valve are more nearly
equal. The young shell (PL XXXVII, fig. 6a) has rather a subrhomboidal shape,
exactl}^ as D. lacteolus, which is due to the better development of the angle between
the upper and posterior margins, which appears as slightly elevated (alate). Thus
the 5'Oung shell is slightly higher in its posterior section, with the posterior end of
the shell less elevated above the base-line, giving to the whole shell a slightly oblique
appearance. But this juvenile shape is sooner or later obliterated, the posterior
wing disappearing, and the posterior end becoming more tapering, giving to the
shell a rather subovate outline, with the posterior end subpointed; but this point
is never very distinct. The outline of Von Ihering’s greeffeanus (PI. 4, fig. 8) comes
very near to the normal shape of the old shell of D. mogymirmi. The brownish
color of the epidermis of our species is also characteristic, and the complete absence
of distinctly greenish tints is to be noted.

Anatomy. — I have a great number of the soft parts of males and of barren and
gravid females, the latter partly with eggs, partly with glochidia in various stages
of development. Of fifty specimens the anatomy has been investigated more
closely. For the breeding season the date of collection (August 28) should be
noted.

Color of soft parts brownish white.

Anal opening slit-like, closed above; closed part not quite twice as long as
the open part. Anal about as long as the branchial, separated from it by a solid
mantle-connection. Branchial opening with distinct papillae. Palpi moderate,
subtriaiigular, posterior margins connected for about one-third of their length.

Gills (PI. XLVI, fig. 5a, b, c) rather wide, the outer one subtriangular, pos-
teriorly slightly wider than the inner; the inner one subtrapezoidal, anteriorly
wider than the outer, its anterior end immediately behind the palpi. Inner lamina
of inner gill entirel}^ connected with abdominal sac. Non-marsupial gills (PI.
XLVI, fig. 5a) with few, scattered interlaminar connections. The marsupiimi
(PI. XLVI, fig. 5b) of the females located in the inner gills, but occupying only
a part, anteriorly leaving free not quite one-third of the gill, and posteriorly hardly
one-fourth, so that the marsupium gravitates slightly toward the posterior section
of the gill. In young females, the marsupial part (PL XLVI, fig. 5b) is much
smaller, and lies distinctly behind the middle of the gill. Structure of the mar-
supium (PI. XLVI, fig. 5c and PI. XLVIII, fig. 2a, b) quite peculiar, consisting of
uninterrupted septa, forming well isolated water-tubes. An interrupted or reticu-

524

MEMOIRS OF THE CARNEGIE MUSEUM.

lated arrangement of the interlaminar connections is nowhere to be seen. Never-
theless this structure must be regarded as developed out of the interrupted con-
dition of the septa, since the non-marsupial gills distinctly show the latter (PI.
XL VI, fig. 5c). In consequence of the development of the water-tubes, the egg-
masses fill these tulies (the ovisacs) in placenta-like bodies (conglutinated) ; how-
ever, these are not very solid and persistent.

(Jlochidium (text-figure 4/v, p. 469) subtriangular, longer than high, rather
small. L. 0.29, II. 0.23 mm. They are slightly oblique, with the lower point
vertically under the posterior third of the hinge-line. They have hooks of the
usual shape, about 0.09 mm. long. Immature glochidia have no hooks.

21. Diplodon suavidicus (Lea) (1856).

Unio suavidicus Lea, Obs., VI, 1857, PI. 29, f. 24.

Diplodon suavidicus Simpson, 1900, p. 876; 1914, p. 1240.

Type-locality. — River Amazon .

New Locality. — Rio Tapajos, Santarem, Para, Brazil (J. D. Haseman coll.,
December 6-12. 1909). Six specimens and three isolated right valves.

Description. — Shell small, greatest L. 28 mm., moderately solid, angularly
subovate or subtrapezoidal, but little oblique, slightly narrower in front, somewhat
broader and sub-pointed behind. Height 69 to 76 pr. ct. of length. Valves not
gaping; dorsal margin gently curved, descending posteriorly, anteriorly descending
more steejily, and passing insensibly into the anterior margin, posteriorly passing in
a rather distinct (indistinct only in largest specimens) but blunt angle into the
posterior margin, which descends obliquely, and is straight or very gently curved.
Lower margin ascending gently in its posterior part, and meeting the posterior
margin in a more or less distinct, but rounded, angle, forming the posterior point
of the shell, which is a little elevated above the base-line. Anterior part of lower
margin longer, sloping distinctly upward. It may be almost straight, or very
gently curved, curving up into the anterior margin. Thus the anterior part of
the shell appears somewhat narrower than the posterior.

Valves moderately convex, slightly flatter on the sides. LTmbonal ridge
rather distinct, but rounded. Posterior slope compressed, produced in young
specimens into a slight wing-like elevation of the jiosterior angle of the upper
margin. In some siiecimens there is a bare indication of a radial rib upon the pos-
terior slope. Diameter 43 to 50 pr. ct. of length. Beaks not much swollen, little
elevated above hinge-line, located at 22 to 29 pr. ct. of length. Beak-sculpture
rather well developed, extending upon the umbonal ridge about 10 mm. or more,

ortmann: south American naiades.

525

but not quite so far upon the rest of the shell, and covering altogether about one-
third or one-fourth of the shell. There are about fifteen or sixteen radial bars, of
which the eighth and ninth unite, and between the latter there is another shorter
pair, also united in v-shape. These bars are rather sharp, those just in front of the
umbonal ridge are hardly broader. The most anterior liars are sometimes slightly
granular, occasioned by the growth-lines cutting across them. There are generally
close to the beaks a few additional radial bars upon the posterior slope, and below
then some oblique, irregular wrinkles, sometimes crossing the former, so as to
form V-shaped angles (forming the posterior system of re-entering angles of Lea).
Lunula present, short, narrower or wider.

Epidermis with numerous, fine, concentric growth-lines, sublamellar on pos-
terior slope, and in the larger specimens with traces of radial lines, chiefly in the
front part of the shell. Color brown to blackish brown, without color-markings.

Hinge-line gently curved. Ligamental sinus over the posterior third of the
laterals. Lateral teeth gently curved, thin, one in right, two in left valve. Pseudo-
cardinals obliquely descending, rather long,' compressed and thin, two in right
valve, the posterior more elevated, with serrated edge. In the left valve also
two pseudocardinals, the anterior larger, serrated, the posterior much smaller,
sometimes rudimentary.

Cavity of shell and beaks moderate. Nacre whitish. Anterior adductor-
scar moderately impressed. Anterior retractor-scar separated, and anterior pro-
tractor-scar united with adductor-scar. Posterior scars indistinct, united. Dorsal
scars in beak-cavity.

Measurements.

No.

Lengtli.

Height.

Diameter.

Beaks.

1

18 mm.

13 mm. = 72 pr. ct. of L.

9 mm. = 50 pr. ct. of L.

at 4 mm. = 22 pr. ct. of L.

18.5 “

14 “ =76

9 “ =49

5 “ =27 “

4

21

15 “ =71

9 “ =43

6 “ =29

5

25

18 “ =72

11.5 “ =46

7 “ =28

5a

27 “

18.5 “ =69

12 “ =44

7.5 “ =28

6

28

20 “ =71

12 “ =43

8 “ =29

Lea’s figure .

21

15 “ =71

9.5 “ =45

5 “ =24

Remarks. My specimens are rather uniform in shape and proportions. There
is some variation in the posterior part of the ventral margin, which may ascend a
little more decidedly, giving to this margin a more distinct lower projection, and
placing the posterior point of the shell a little higher above the base-line. In
my largest specimen, all angles (upper iiosterior, iiosterior, and that of lower
margin) are more rounded. The beak-sculpture may be more or less distinct and
sharp, and the bars vary somewhat in length.

526

MEMOIRS OP THE CARNEGIE MUSEUM.

Previously this species was known only from a single individual described
by Lea. There is not the slightest doubt that my specimens belong here, and one
of them (No. 4) is almost a replica of that of Lea. Lea believes that his shell is
a young one, and it certainly is not full-grown, but, as my material shows, this
species does not grow very much larger. Von Ihering (1893, p. 120) suggests
that this might be the young stage of U. wheatleyanus Lea, but this cannot be the
case, since the latter has a different, much heavier beak-sculpture.

I am uncertain about the systematic position of this species, but there are
certain resemblances in the shape of the shell to that of the lacteolus-group. This
species is .also interesting because of the fact that it is one of the few forms of
Diplodon found in the Amazon-drainage.

G. Group of Diplodon ellipticus.

Shell more or less elongated, but often rather high and short; subovate or
siibtrapezoidal, distinctly oblique at all stages of growth, with the longest axis
forming an angle with the line of the ligament. Anterior end narrower, posterior
end higher, and lower margin distinctly ascending in its anterior portion. Beak-
sculpture fine or coarse, more or less developed.

The chief character of this group is the obliquity of the shell, brought about
by a widening of the posterior portion of the shell in the vertical direction, so that
the posterior end lies rather low, and the longest dimension is not parallel or nearly
parallel to the ligament, but forms a distinct angle with it. Although an obliquity
is sometimes indicated in the species of the previous groups, it generally disappears
with increasing age, while in the present group the obliquity is rather emphasized
in older shells. It is all-important, in order to correctly judge as to the shape of
these shells, to place them in a uniform position, always with the ligament running
horizontally.

The outline of the shells of this group varies a good deal, and some of them
become very high in proportion to length, so that the outline appears more nearly
subrotund, similar to the shape in the subgenus Cycloniya. However, in the
latter the greatest height of the shell is always situated more nearly in the middle
under the middle of the ligament, while in the shells of the ellipticus-group the
greatest lieight is more posteriorly, at the posterior end of the ligament, or even
beyond that.

22. Diplodon ellipticus Spix (1827).

Diplodon ellypticmn (error typogr.) Spix, 1827, PI. 26, fig. 1-2.

Unio ellipticus Wagner, 1827, p. 33; Von Ihering, 1890, p. 163, PI. 9, figs. 8-9;

Von Ihering, 1893, p. 108.

ortmann: south American naiades.

527

Diplodon wagnerianum Simpson, 1900, p. 877; 1914, p. 1246.

Type-locality. Rio San Francisco (Wagner).

Other Localities. — Rio Parahyba do Sul, Rio de Janeiro (Von Ihering, 1893, p.
115); Rio Santa Maria, Espirito Santo (Von Ihering, 1910, p. 134) (drainage of
Rio Doce). The form from the latter locality has been named var. santanus Von
Ihering.

Rio Piracicaba, Sao Paulo, and^Rio Tamanduatahy, Sao Paulo (Von Ihering).
The latter two localities are somewhat doubtful, since the specimens are not exactly
like ellipticus. The location and drainage of the last named river is unknown to
me.

In the Carnegie Museum is one specimen, labeled “Brazil” (Holland Collec-
tion). Not quite typical.

The change of the specific name ellipticus to wagnerianus is unwarranted.
Spix gave the name in the plate in connection with the generic name Diplodon.
Since the latter stands, the specific name also is entitled to recognition.

In spite of Von Ihering’s re-description of the type, this species is as yet poorly
known, and our knowledge of it is founded chiefly upon what Von Ihering has
said. The species is positively known from the Rio San Francisco and the Rio
Parahyba do Sul; the other localities are more or less doubtful, since the specimens
described from them do not fully agree with the type. Those from the Rio Santa
Maria have been distinguished as a variety.

All we can gather from descriptions and figures is that D. ellipticus is a sub-
elliptical or subtrapezoidal shell, of dark green to blackish color, with rather smooth
surface. It has in the anterior part of the shell a shallow depression, often producing
a shallow emargination in the anterior part of the ventral margin. In outline,
the shell is distinctly oblique, narrower in front, higher behind. The beak-sculpture
consists of simple, fine radial bars, which are rather short, and somewhat cut up
on the posterior slope by irregular, oblique wrinkles. The nacre is blueish white.
Pseudocardinals somewhat compressed, but not thin, crenulated, two in right,
one in left valve, but the latter with an angle at its base, representing the remnant
of a posterior tooth.

Our specimen from Brazil was received as ellipticus. It agrees in most of the
above characters, except that it is shorter in proportion to height, with less pointed
posterior end. Also the projection of the lower margin is indistinct. The beak
has eleven radial bars in front of the posterior ridge, the seventh and eighth meeting
in the middle. On the posterior slope there are corrugations and fine, oblique
ridges. L. 32 mm.; H. 21 mm.; D. 12 mm. This specimen might very well be

528

MEMOIRS OF THE CARNEGIE MUSEUM.

R young ellipticus, and may correspond to the var. santanus of Von Iliering (smaller,
l)rojection of lower margin less distinct).

23. Diplodon berths Ortmann, sp. nov.

Shells: Plate XXXVIII, figs. 1, 2, 3, 4; Anatomy of gills: Plate XLVI, fig. 6.

Type-locality . — Rio Jacuhy, Cachoeira, Rio Grande do Sul, Brazil (J. D. Hase-
man coll., January 2G, 1909). Type-set: Cam. Mus. Cat. No. 61.58G5. Sixteen
sjjecimens, all with soft parts, including males, barren and gravid females. (There
were twenty-three specimens in the original lot).

Additional Loeality. — Rio Vaccahy Mirim, Santa IMaria, Rio Grande do Sul,
Brazil (J. D. Ilaseman coll., January 29. 1909). One male with soft parts.

Distribution: Cuahyba drainage in southern Brazil.

Description of Shell. — Shell rather small, maximum length G5 mm.; rather
solid, chiefly so anteriorly, and in old shells often much thickened along the lower
anterior margin. Outline subovate to subtrapezoidal, distinctly oblique, broad
and rounded, or somewhat jiointed behind. Height from 55 to G7 iir. ct. of length.
Valves not gaping. Dorsal margin straight, or gently descending posteriorly,
forming a more or less distinct obtuse angle with the jiosterior margin. The latter
obliquely descending, gently convex, and curving around into the posterior part
of the lower margin, forming with the latter in young specimens an indistinct
rounded angle, which, however, may become more distinct in old specimens. Lower
margin in normal siiecimens with a distinct rounded iirojection, forming the lowest
])oint of this margin, situated far back, behind the posterior end of the ligament.
From this ])oint the lower margin curves up behind to the posterior end of the shell
and this part is quite short. Anteriorly the lower margin also slopes upward,
and is almost straight for a considerable, distance; sometimes it is even slightly
concave; then it curves up into the anterior margin. Thus the shell appears con-
siderably narrower anteriorly, broader (higher) iiosteriorly, with the greatest
height situated far backward.

V alves moderately and not uniformly convex. The greatest convexity is near
the anterior end and over the iiosterior ridge, which is broad and not sharply marked.
In front of the posterior ridge the sides of the disk are distinctly and broadly flat-
tened, and sometimes even slightly concave, producing the emargination of the
anterior part of the lower margin. Posterior slope somewhat compressed, very
rarely with a slight trace of a rib or a furrow. Greatest diameter of the shell 32
to 43 })r. ct. of the length, located well behind, upon, or close in front of, the pos-
terior ridge. Thus, although rather swollen in the region of the posterior ridge.

ortmann: south American naiades.

529

the shell appears in front of it rather more compressed. Beaks not much swollen,
and not very prominent, located at from 23 to 29 i)r. ct. of the length. Beak-
sculpture consists of twelve to fourteen radial bars, which are sharp and rather
distant from each other, the ninth, tenth, and eleventh (immediately in front of
and upon the posterior ridge) are longest, about 10 to 12 mm. long; the seventh
and eighth may consist of two bars united in V-shape, l)ut this cannot be seen
clearly, since in all specimens the tips of the beaks are eroded. These median
bars (seventh and eighth) are also slightly shorter than those in front, and distinctly
shorter than those behind them; the last two or three bars are fine and somewhat
shorter, and stand upon the posterior slojie, becoming indistinct. In a few of my
specimens there are traces of oblique wrinkles upon the posterior sloiie. Lunula
short and narrow, distinct only in larger specimens.

Epidermis smooth, rather shining, with numerous, closely set, fine, concentric
lines, and stronger and irregular concentric wrinkles. The fine lines become sub-
lamellar on the posterior slope and towards the margins. A fine radial sculpture
is present, chiefly in the anterior iiart of the shell, but it is not very evident. Color
greenish black to brownish black, darkest in old shells. Young shells are more
distinctly greenish, dark olive-green, shading towards the beaks to gray-green and
brownish olive. There are no distinct color-bands and no distinct color-rays,
except in very young specimens, where there are traces of dark green rays on the
posterior slope, seen when held iqi against a strong light.

Hinge straight or gently curved. Ligamental sinus over the ]iosterior third
or fourth of the laterals (more iiosteriorlj^ in old shells). I^ateral teeth curved,
more strongly so in their posterior part in old shells, one in right, two in left valve,
edges rugose. Pseudocardinals two in right, one in left valve, subcompressed,
not very long, in old siiecimens sometimes almost stumpy. The posterior })seudo-
cardinal of right valve stronger and more elevated than the anterior, often much
divided, and always much crenulated. Very often there is a second posterior
small pseudocardinal in the left valve.

Cavity of shell and beaks shallow. Nacre whitish, iri old specimens often
very thick anteriorly, and with pinkish tints; posteriorly very iridescent. Ad-
ductor-scars deeply impressed anteriorly, less so posteriorly. Anterior retractor-
scar small and deep, separated from the adductor-scar; anterior protractor-scar
united with it. Posterior retractor-scar connected with adductor-scar. Pallial
line distinct. Dorsal scars in an oblique irregular row in the beak-cavity.

Remarks. — This species may be what Von Ihering (1893, p. 102) calls athiops
in the Guahyba drainage, but only the remark (]). 104) that this aiJiiops has a

530

MEMOIRS OF THE CARNEGIE MUSEUM.

broad, shallow furrow in the anterior part of the shell, seems to confirm this as-
sumption. In other respects it is impossible to decide, whether this is, or is not, the
ccthiops of Von Ihering. It surely is 7wt the oethiops of Lea. Of the few other
species incidentally mentioned by Von Ihering as found in the Giiahyba-drainage,
none can be compared with onr species. Its chief characters are the subovate to
snbtrapezoidal distinctly oblicpie shape, narrow in fiunt, broader behind, and the
peculiar compression of the shell in the anterior part. The beak-sciilptiire and
color of the epidermis are also characteristic.

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Figured.

3.. .

n

&

rpe-set

31.5 mm.

21 mm

= 67 pr. ct. of L.

10

mm. =32 pr.

’t. of L.

at 8

mm. =25 pr. ct. of L.

PL XXXVIII, fig. 2.

11.. .

cf

43.5 “

27.5 “

= 63

18

“ =41

12.5

“ =29

14.. .

9

47

31.5 “

= 67

19

“ =40

12.5

“ =27

22.

9

51.5 “

32

= 62

22

“ =43

12

“ =23

23.. .

9

52

29

= 56

21.5

“ =41

12

“ =23

24. . .

c?

56

36

= 64

21.5

“ =38

13.5

“ =24

PI. XXXVIII, fig. 1.

.34. . .

cT

65

35.5 “

= 55

26

“. =40

15

“ =23

(Santa Maria)

I cannot compare this species with any other, except D. ellipticus Spix. The
general shape is very much the same, but ellipticus seems to be more elongate
(height only 54 pr. ct. of length). The jiosterior end is slightly more pointed, and
the diameter is distinctly less (31 iir. ct. on the average). In our species the average
height is 62 pr. ct. and the diameter 39 pr. ct. In addition our shell seems to be
thicker and more solid, chiefly in old specimens, where the lower anterior margin
is considerably and strikingly thickened, a character not mentioned in ellipticus.

Old shells often become freakish, assuming irregular shapes. Frequently the
liosterior part of the shell grows more strongly in length, thus rendering the shell
exceptionally long (as in No. 23). In such specimens the projection of the lower
margin is obscured, and the posterior point of the shell is very little elevated above
the base-line. Furthermore the shell in general is more pointed behind. Such
siiecimens also ajipear more swollen. However, the growth-lines clearly indicate
that, when young, these individuals had the normal shape. In other old shells
the whole posterior part is more developed (No. 24, PI. XXXVIII, fig. la) and is
deflected downward. This fact tends to preserve the general shape, but renders the
anterior ])art of the lower margin somewhat concave.

Anato7ny. — Judging from the soft parts at hand, twelve specimens are males,
five are barren females, and seven are gravid females. Only one of the latter con-
tained immature glochidia. The date of collection (January 26) apparently is
near the beginning of the breeding season.

ortmann: south American naiades.

531

Color. — Distal part of foot dark brownish, grayish, or lilackish; the rest of
the soft parts are whitish.

Anal opening closed above ; the closed part about four times as long, or a little
longer than the open part; the latter slit-like, slightly shorter than the branchial
opening. The latter with small, but distinct papillie, separated from the anal by
a solid mantle-union. Palpi subtriangular, moderately large, of the usual shape;
their posterior margins connected at base.

Gills rather long and wide. Outer gill subtriangular, widest at the beginning
of the posterior third, and here it jirojects a little beyond the inner gill. Inner gill
subtrapezoidal, its anterior end immediately behind the iialpi. Inner lamina of
inner gill connected with abdominal sac.

Structure of non-marsupial gills as usual. In the female, the marsupium (PI.
XLVI, fig. 6) is located in the inner gill, and in large specimens it is restricted to
the middle part of the gill, leaving non-marsupial almost one-third at the anterior
end, and a somewhat smaller portion at the posterior end. In young specimens
the marsupium is smaller. When charged, the marsupium forms a slightly swollen,
lenticular, rounded, or oblong patch in the middle of the gill, most of it lying behind
the middle. The interlaminar connections of the marsujiium are strongly de-
veloped, forming very incom})lete, interrupted sejita, and arranging themselves
rather in transverse and obliipie rows, so that the vertical se])tiform structure is
obscure, while a reticulate and irregularly quincuncial arrangement prevails.
Only near the margin of the gill is a septiform structure indistinctly indicated.
The egg-masses do not conglutinate into jilacentin-like structures.

Only one of my females has very young glochidia. They are, as far as can be
seen, of the usual shape, subtriangular, and somewhat oblique. Exact measure-
ments could not be obtained. No hooks are visible, but, of course, such may be
present in ripe glochidia.

24. Diplodon enno Ortmann, sp. nov.

Shells: Plate XXXVIII, figs. 5, 6, 7, 8; Anatomy of gills: Plate XLVI, fig. 7.

Type-locality. — Rio Grande, Boqueirao, Bahia, Brazil (S. Francisco drainage).
(J. D. Haseman coll., January 9, 1908). Type-set: Cam. Mus. Cat. No. 31.9264.
Eighteen specimens, males and barren females, with soft parts. (A number
of additional young specimens were in the original set.)

According to the latest census of the Naiades from the Rio S. Francisco
drainage (Von Ihering, 1910, p. 138), there are only two species of Diplodon present
•in this system; D. rotundus Spix, and D. ellipticus Spix. The former is much

532

MEMOIRS OF THE CARNEGIE MUSEUM.

higher and much more rounded than the present siiecies; the latter is more elongated
and more jiointed behind, and has, besides, a smooth epidermis {Cf. Von Ihering,
1890, p. 163). Since it is imjiossible for me to find any other South American
species of Diplodon, the description of which answers to the present species, we
must regard the latter as new.

Description of Shell. — Shell small to medium (maximum length 53 mm.),
rather thin. Outline subovate or subtrapezoidal, distinctl}^ oblique, higher behind,
narrowed anteriorly, the obliquity being most pronounced in older specimens.
Height 56 to 75 pr. ct. of length (against 54 pr. ct. in ellipticus, and 86 or 87 pr. ct.
in rotundus). Valves not gaping. Dorsal margin straight or gently convex, form-
ing a rather distinct or obtuse angle with the posterior margin, which descends ob-
liquely and rather steeply, is nearly straight or gently convex, and curves into
the lower margin without forming a distinct iiosterior point. The posterior ex-
tremity of the shell is located relatively low and is only moderately elevated above
the base-line. Lower margin with its lowest point located rather posteriorly,
vertically below the jiosterior end of the ligament, or even behind it, strongly as-
cending in a curve toward the posterior end of the shell, but nearly straight or
very slightly convex in its anterior part, and sloping upward toward the anterior
margin, so that the anterior ]iortion of the shell is distinctly narrower than the
posterior, producing thus the oblique aiipearance of the whole shell.

Valves com]iaratively comjiressed. Diameter 28 to 45 pr. ct. of length.
Greatest convexity and greatest diameter situated well back upon the posterior
ridge, which, however, is very indistinct and broad. Sides of the shell in front of
posterior ridge not very convex and rather flattened. Posterior slope compressed,
sometimes with a faint trace of a radial groove, somewhat elevated (wing-like)
toward the up])er jiosterior angle. Beaks not swollen, and hardly elevated above
the hinge-line, located at from 22 to 28 pr. ct. of the length. Beak-sculpture
sharp and fine, distinct, but restricted to the region near the beaks. There are
fifteen to eighteen radial bars, the ninth and tenth in the middle, joined and v-
shajied at their lower ends. The longest bars (6 to 8 mm.) stand upon the posterior
ridge, the anterior ones being distinctly shorter. There are a few anterior bars
and iqion the iiosterior slojie a few posterior bars, and sometimes traces of oblique
wrinkles. None of the bars are distinctly granular, and a comparatively slight
degree of erosion obliterates all traces of beak-sculpture. Lunula indistinct and
narrow, visible onl^^ in older shells.

Ll)idermis not shining, but rather rough. This is due to a great number of
fine, irregulai“, concentric lines, which, when well-preserved, are lamellar and

ortmann: south American naiades.

533

elevated, showing this character all over the disk, being, however, more distinctly
lamellar and more crowded upon the posterior slope and near the margins. Even
when these fine lamellae are worn off, the epidermis does not become shining, but
remains dull, and when well-preserved, the epidermis appears cloth-like. There
are no traces of radial sculpture. Color of epidermis dark greenish black, but
often in the middle of the disk and towards the beaks brownish black. The greenish
tint is not very evident, and is best seen in young specimens.

Hinge-line very gently curved. Ligamental sinus over the posterior third of
the laterals. Lateral teeth curved, one in right, two in left valve, their edges
slightly corrugated or nearly smooth. Pseudocarclinals narrow and compressed,
but not very long, corrugated and rugose, but not cut u}}, two well developed
pseudocardinals of nearly equal size in right valve, one well-developed in left valve;
but. there often is a smaller posterior one in the left valve, and another small, low,
and narrow anterior one, so that the left valve may have three pseudocardinals.
The middle one, however, is always the largest. Sometimes the posterior pseudo-
cardinal of the right valve is higher and thicker than the anterior.

Cavity of shell and beaks shallow. Nacre blueish silvery, very iridescent,
and sometimes discolored yellowish. Anterior adductor-scar distinct and moder-
ately impressed. Anterior retractor-scar rounded, small and rather deep, separ-
ated from adductor-scar. Anterior protractor-scar united with it. Posterior
adductor-scar rather indistinct and not impressed, the posterior retractor-scar
forming an upper process of it. Pallial line not very sharp. Dorsal scars few in
beak-cavity.

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Figured.

18.. .

?

12.5 mm.

7 mm

=56 pr.

ct. of L.

3.5

mm. =28 pr. ct. of L.

at 3.5

mm. =28 pr.

ct. of L.

14.. .

?

22

13.5 “

=61

7

“ =32

6

“ =27

8.. .

28

20.5 “

=73

9

“ =32

7

“ =25

4.. .

d'

35

26

=74

12

“ =34

8

“ =23

PI. XXXVIII, fig. 5.

2.. .

9

45

32

=71

16.5

“ =37

10

“ =22

PI. XXXVIII, fig. 8.

1.. .

9

53

40

=75

24

“ =45

12.5

“ =24

PI. XXXVIII, fig. 7.

Remarks. — The dimensions of this species change slightly with age, in that
younger shells are not so high in proportion, and consequently, they appear
less distinctly oblique than older ones. The diameter of young shells is also less
than in old ones, and the highest figures (37 to 45 pr. ct.) are shown by old females.
In other respects my specimens are rather uniform, and the oblique, not very
elongated shape, and the dull, blackish epidermis and silvery nacre are chiefiy
characteristic.

534

MEMOIRS OF THE CARNEGIE MUSEUM.

Anatomy. — It was not possible to positively ascertain the sex of my smallest
specimens. But, I have other specimens, of which five are positively males, and
seven of which are females. None of the latter were gravid.

Color of soft parts whitish, distal part of foot brown.

Anal opening closed above; closed part considerably longer than the open
part; the latter slit-like, shorter than the branchial opening, and separated from
it by a solid mantle-connection. Branchial opening with distinct papillae. Palpi
siibtriangular, their posterior margins connected for about one-third of their
length.

Gills of the usual shajie, rather wide, the inner anteriorly wider, its anterior
end close to the paljii. Inner lamina of inner gill entirelj^ connected with abdominal
sac. Non-marsupial gills of the usual structure, with few and scattered inter-
laminar connections. Marsupium of the female (PI. XLVI, fig. 76) located in
the inner gill, occu])ying nearly half of it in the middle, leaving non-marsupial
one-fourth of the gill l^oth at the anterior and posterior end. In young females
the marsupial jiart is smaller (PI. XLVI, fig. 7a), but also located nearly centrally.
Interlaminar connections of the marsupium reticulated or cpiincuncial toward the
base, but forming interrupted septa toward the margin.

25. Diplodon gratus (Lea) (1860).

Section of gills: Plate XLVIII, fig. 3.

Unio grains Lea, Obs., X, 1863, PI. 43, fig. 290; Sowerby, XVI, 1868, PI. 84, fig. 444.
Diplodon {Cyclo7nya) grains Simpson, 1900, p. 886.

Diplodon (Cyclomya) fontainianus gratus Simpson, 1914, p. 1281.

Type-locality. — Rio Uruguay.

New Locality. — Rio Uruguay (in mud), Uruguayana, Rio Grande do Sul,
Brazil (J. D. Haseman coll., February 5, 1909) twenty-one specimens, males
and females, all with soft parts.

Distribidion. — Positively known only from the Uruguay River. Von Ihering
(1893, p. 92) gives this form also from Rio Guahyba, Porto Alegre, Rio Grande
do Sul. However, as we shall see, this is not the typical grains, and the Guahyba-
form has already been distinguished by Simpson as var. deceptus (see below).

Description of Shell. — Rather small, maximum length of my specimens 53 mm.,
moderately thick. Outline obliquely subrotund, subovate, or subtrapezoidal,
very variable, more or less high (height 72 to 85 pr. ct. of length). There is an
indistinct relation of the shape to sex, in so far that the highest shells (79 pr. ct.
and over in height) are all males, while the more elongated specimens may be either

ortmann: south American naiades.

535

males or females, the former being more subrotund, the latter more subovate-
subtrapezoidal. Dorsal margin gently convex, forming a more or less distinct,
obtuse angle with the posterior margin, the latter obliquely descending, straight,
or gently curved, curving broadly around into the lower margin, forming the in-
distinctly defined, rounded, posterior end of the shell. Lower margin ascending in
its anterior part, gently curved in the higher shells, or almost straight in those
more elongated, its posterior part curving up towards the posterior margin. The
lowest point of the lower margin (and greatest height of shell) is located well back-
ward, vertically below the posterior end of the ligament, or even behind this. Thus
the shell presents a rather oblique shape, being narrowed anteriorly, and higher
posteriorly, but, according to the varying height, this obliquity is more or less
pronounced, most distinctly in the more elongated shells, less so in those more
elevated, where the outline approaches the subrotund shape.

Valves moderately but variaJjly convex, convexity greatest upon the broad
posterior ridge, smallest upon the sides of the disk, where elongated specimens
are almost flat. Posterior slope slightly compressed, sometimes with a trace of a
radial furrow and a ridge, which, however, may be entirely obliterated. Diameter
36 to 50 pr. ct. of length. The more compressed specimens are the smaller, but
no distinct relation to sex can be discovered in this. Beaks moderately swollen,
not very prominent, located at 25 to 34 pr. ct. of the length (more anteriorly in
larger specimens). Beak-sculpture weakly developed, seen only in smaller speci-
mens, consisting of twelve to thirteen radial bars, of which the seventh and eighth
unite v-shapedly. The bai’s do not differ much in length, although the posterior
ones are slightly longer, the maximum being hardly over 8 mm. long. They are
not very sharp, but rather blunt, only the most anterior and posterior (upon pos-
terior slope) are somewhat finer and sharper. Often there are upon the posterior
slope irregular oblique wrinkles, which may be rather numerous, or may be al-
together absent. Lunula absent in young, but sometimes seen in older specimens,
short and very narrow.

Epidermis smooth in the middle of the disk, but not very shining, covered
with numerous somewhat irregular, fine, concentric lines, and more widely separated
irregular concentric wriidvles. The concentric lines are sublamellar iqion the an-
terior end, the posterior slojie, and towards the margin, and sometimes they are
sublamellar all over the disk, but the latter is the case only in young individuals.
In old specimens they undoubtedly have been worn off. Radial sculpture generally
present. It may be very obscure, chiefly in younger shells, but is visible in older
shells, consisting of faint, blunt radial ridges, which are rather irregular, often in-

536

MEMOIRS OF THE CARNEGIE MUSEUM.

terrupted, and more distinct in the anterior part of the shell. Even when best
developed, they do not constitute a prominent feature of the surface. Color of
epidermis dark olive-green to blackish. In young specimens it is more grayish
or brownish green towards the beaks; in older specimens it appears nearly black.
When slightly worn, it is in places lighter, brownish olive, or greenish olive. In some
older specimens brownish tints appear towards the lower margin, and sometimes
a faint brownish concentric band is indicated. The general impression of the
color of the epidermis, however, is blackish green, not brown. No color-rays are
visible.

Hinge-line gently curved. Ligamental sinus over the posterior third of the
laterals, in young specimens over the posterior half. Lateral teeth curved more
strongly in their posterior part in old shells, one in right, two in left valve, their
edges rough. Pseudocardinals directed oblicpiely forwards and downwards,
lamellar and compressed, but not very long. In the right valve there are two of
them, the posterior much larger than the anterior. In the left valve there is one
pseudocardinal, but quite often a smaller posterior one is present. All pseudo-
cardinals are rough, crenulated, or serrated.

Cavity of shell and beaks moderate. Nacre whitish, iridescent, in young
shells blueish or greenish silvery towards the edges. In older shells it becomes
more opaque white, and assumes generally a creamy or salmon hue in and near
the cavity. In a few specimens there is a very delicate pinkish tint all over the
nacre.

Anterior adductor-scar rather deep, subelliptical; anterior retractor-scar
small, deep, isolated from the adductor-scar; anterior protractor-scar connected
with it. Posterior adductor-scar subtriangular, faintly impressed, posterior re-
tractor-scar forming an upper triangular process of it. Pallial line distinct. Dorsal
scars few, in an irregular, oblique line in the beak-cavity.

Measurements.

No.

Se.x.

Length.

Height.

Diameter.

Beaks.

6. . .

?

23.. 5 mill.

17

mm. = 72 iir.

ct. of L.

8.5

mm

. =36 pr. ct. of L.

at 7.5

mm. = 32 pr.

ct. of L.

2 . . .

?

28.5 “

23.5

“ =85

< i

11

U

= 39

9

= 32

( (

3. . .

?

32

23.5

“ =73

i i

14

U

= 44

11

= 34

i (

7 . . .

9

36

26

“ =72

U

13.5

u

= 38

11

i (

= 31

((

13. . .

37.5 “

32

“ =85

n

16.5

( (

= 44

10

((

= 27

U

10. . .

cf

39'

2!)

“ =74

((

16

u

= 41

11.5

U

= 29

a

16. . .

cf

40

32.5

“ =81

((

20

u

= 50

11.5

U

= 29

Li

14. . .

9

41

32

“ =78

( (

18

u

= 44

12.5

u

= 30

Li

22 . . .

47

39.5

“ =84

20

u

= 43

12

iC

= 26

ii

21 . . .

9

.50

36

“ =72

((

21

u

= 42

14

it

= 28

ii

Half shell..

53

38

“ =72

22

= 42

13

u

= 25

ii

ortmann: south American naiades.

537

Remarks. — According to Simpson (1914), this form (gratiis) is a variety of
D. fontainianus (D’Orbigny), and there is no doubt that both are closely allied to
each other. However, Simpson erroneously gives the distribution of fontainianus
as: ‘‘Uruguay River and its affluents; Parana River, southern Brazil,” while it
is known only, according to D’Orbigny and Von Ihering (1893) from the Rio
Parahyba, Rio de Janeiro, the upper Parana-drainage in Sao Paulo, and Lagoa
Santa in Minas Geraes (Rio de las Velhas, Sao Francisco-drainage) . In 1910 (p.
138) Von Ihering drops this latter locality. Fontainianus has never been rejDorted
from the Uruguay. Thus it is not very likely that my specimens from this river
are D. fontainianus, although they agree with it in some particulars.

According to Von Ihering’s account of D. fontainianus (1893, p. 90, PI. 4, fig. 6),
the latter differs from my specimens chiefly in having the posterior retractor-
scar separated from the adductor-scar, but otherwise they are very similar. This
holds good chiefly in the color of the epidermis. My specimens are greenish black,
which agrees best with Von Ihering’s D. fontainianus, which is “black, with greenish
basis,” and to a degree with D’Orbigny’s “brim noiratre,” while Lea describes his
gratus as “dark olive brown.” The differences between D. gratus and D. fontaini-
anus, as given by Simpson, concern the shape of the shell, which is said in gratus
to be more wedge-shaped in front, and the texture of the epidermis, which is said
to be smoother, subshining, and lighter in color. These differences I cannot
recognize in my specimens. It is therefore hard to decide by which name they
should be called. However, my specimens coming from the Uruguay River,
being practically topotypes of gratus, I have concluded to call by this name, and,
in order to avoid any misunderstanding, I have given a full description of them.

Von Ihering (1893) believes that he has discovered sexual differences in the
shell of D. fontainianus, the males being higher and shorter, and more rounded,
the females being longer and more sub trapezoidal. Similar differences in shape
are also noticed in our D. gratus. I have seventeen specimens in which the sex has
been positively ascertained by examination of the gills, and I find that it is not
possible to accurately determine the sex according to the shape of the shell. First
of all, the two shapes are not sharply separated, but pass insensibly into each other,
many specimens being intermediate between the two extremes. Besides, if form
has any relation to sex, we may say that the highest shells (with the height of 79
pr. ct. of length and over) are, indeed, males; but among the others, having the
height of 72 to 78 pr. ct. of length, there are as many females as males, and this ap-
plies both to larger and smaller shells. It is true, that the average height of all my
males is about 78 pr. ct., and the average height of all my females is 75 pr. ct.; but

538

MEMOIRS OF THE CARNEGIE MUSEUM.

these differences are so slight, that they do not furnish an exact criterion for the
distinction of the sexes.

Aiiatomif-^. — The soft parts of eleven males and six females have been in-
vestigated.

Color of distal part of foot blackish, this color sharply set off from the pale
basal part. Rest of soft jiarts whitish. Outer edge of anal opening (including
the closed part) and of branchial, black; this color running forward beyond the
branchial opening for a short distance.

Anal opening closed above; closed part much longer than the open, the latter
slit-like, a little shorter than the branchial opening, and separated from it by a
solid mantle-connection. Branchial opening with small, but distinct papillae.
Palpi rather large for the size of the species, subtriangular, with curved lower
margin; jiosterior margins connected only at base.

Gills moderately long and rather wide. Outer gill with strongly convex lower
edge, its widest part a little back of the middle, and about as wide as the inner
gill in its posterior jiart. Anteriorly it becomes narrower, and its anterior end is
near the highest point of the mantle-attachment-line. Inner gill with the inner
lamina entirely connected with abdominal sac, its edge nearly straight in the an-
terior part, where it is much wider than the outer gill. Anteriorly it is slightly
narrowed, and its anterior end is immediately behind the palpi. Structure of
non-marsupial gills (PI. XLVIII, fig. 3a) somewhat unusual, with rather crowded,
irregular interlaminar connections, forming weak, interrupted septa; these con-
nections are more frequent toward the margins of the gills. Inner gill of the fe-
male marsupial. Its interlaminar connections (PI. XLVIII, fig. 36) are more
crowded, and heavier, arranged in interrupted rows running parallel to the gill-
filaments, forming incomplete septa. The marsupium occupies only a part of the
gill, leaving a small section at the anterior end free, a somewhat larger one (but
hardly one-fourth) at the iiosterior end free also, so that the marsupium is located
in the middle of the gill, and slightly more anteriorly than posteriorly.

In this species, the structure of the marsupial and non-marsupial gills is more
alike than usual as regards the arrangement and frequency of the interlaminar
connections, but in the marsupium the connections are decidedly heavier and
stronger, and form more distinct, although interrupted, septa (See PI. XLVIII,
figs. 3a and 36).

The anatomy of D. foniainianus has been discussed by Von Ihering (IS93, p. 90), but the finer
structure of tlie marsupium has not been describeil. The glocliidia are said to be subtriangular, without
hooks. Their length is 0.4 to 0.5 mm. Tlius they are unusually large for the genus, and belong to the
largest known among all Naiades.

ortmann: south American naiades.

539

26. Diplodon deceptus (Simpson) (1914)

Shells: Plate XXXIX, figs. 1, 2, 3, 4, 5; A7iatoniy of gills: Plate XLVII, fig. 1;

Section of gills: Plate XLVIII, fig. 4.

Unio grains Von Ihering {non Lea), 1893, p. 92.

Diplodon {Cyclornya) fontainianus deceptus Simpson, 1914, p. 1281.

Type-locality . — GuahjLa, Brazil.

Other Locality. — Rio Giiahyba, Porto Alegre, Rio Grande do Sul, Brazil (Von
Ihering) .

Localities Represented in Carnegie Museum. — Rio Guahyba, Porto Alegre,
Rio Grande do Sul, Brazil (J. D. Haseman coll., January 24, 1909) two shells
and one right valve {Topotypes). Rio Jaciihy, Cachoeira, Rio Grande do Sul,
Brazil (J. D. Haseman coll., January 26, 1909) ten specimens with soft parts,
males and females.

Distribution. — Drainage of the Rio Guahyba and its tributary, the Rio Jacuhy,
in Rio Grande do Sul.

There is no question that my specimens belong to the form called by Von
Ihering grains, and by Simpson deceptus. Von Ihering compares his D. grains
with D. fontainianus, and says that the former much resembles the latter in shape,
but that the ejiidermis of D. grains is brown, shows radial sculpture, and is smoother;
that the beaks are more anterior (24 to 31 pr. ct., average 27 pr. ct., against 29 to
39 pr. ct., average 34 pr. ct. in D. fontainianus); that the pseudocardinals of D.
grains are shorter and more stumpy; and that the posterior retractor-scar is always
connected with the adductor-scar.

All these characters hold good for my s])ecimens, and they distinguish this
form also from the real D. grains of the Uruguay River, except that the location
of the beaks does not differ so much (in true D. grains it is 25 to 34 pr. ct.).

Simpson’s D. deceptus has been described as being unevenly obovate, subin-
flated, rather solid; pseudocardinals shorter, more stumpy, and much split. This
also fits the present form, and I have no doubt that D. deceptus is the same shell
as that called D. grains by Von Ihering.

The chief differences between D. deceptus and D. grains are the following:

1. D. deceptus has a greater tendency toward the obliquely elongated shape,
with subtrapezoidal or subovate outline. The height varies from 65 to 78 pr. ct.
of the length, while in D. gratus it ranges from 72 to 85 jir. ct.

2. The epidermis of D. deceptus is always brownish, with hardly any green in
it, and not blackish green. It is smoother, since the fine concentric lines are not

540

MEMOIRS OF THE CARNEGIE MUSEUM.

lamellarly elevated (except indistinctly so near the margins), and the radial sculp-
ture is more distinct, consisting of faint furrows, becoming often subscalariform
toward the margins.

3. The pseudocardinals of D. deceptus are more stumpy, chiefly in larger
specimens. In younger ones this difference is not so marked, yet noticeable, when
individuals of the same size are compared. In large specimens they are short
and thick, chiefly so the posterior one of the right valve, and the anterior of the
left (PI. XXXIX, figs. Ic, Id). The posterior one of the left valve generally is
well developed, although smaller than the anterior one. All these teeth are split
up into a number of denticles, and there are frequently accessory teeth, anteriorly
in the left, and posteriorly in the right valve.

All other characters are as in D. grains, but D. deceptus grows to a larger size.

Measukements.

No.

vSex.

Length.

Height.

. Diameter.

Beaks.

Figured.

Gael

loeira

1.. .

?

28 mm.

19 mm

= 68 pr. ct. of L.

9

mm. =32 pr. ct. of L.

at 8 mm

= 29 pr. ct. of L.

4.. .

d'

43

33

=77

18.5

“ =43

13 “

=30

PI. XXXIX, fig. 3.

6.. .

9

47.5 “

35

=74

19

“ =40

15 “

=31

PI. XXXIX, fig. 4.

8.. .

d'

56

39.5 “

=71

22.5

. “ =40

14 “

= 25

11.. .

d

62

40

= 65

27.5

“ =44

17 “

= 27

PI. XXXIX, fig. 2.

9.. .

9

64

43

=67

26

“ =41

17 “

=27

Porto Alegre

2.

49

37

= 76

18.5

“ =38

12 “

=24

4.. .

55

43

=78

26

“ =47

13 “

= 24

Remarks. — In this species we also observe higher and more elongated shells,
nevertheless no relation between shape and sex is apparent. In fact, the most
elongated specimen No. 11, H. 65 pr. ct. (See PI. XXXIX, fig. 2) is a male, and not,
as might be expected (according to Von Ihering) a female. Altogether the number
of individuals with the sex positively known is rather small, but the gradual transi-
tion of the more elongated into the shorter and higher ones is evident.

Anatomy. — From Cachoeira, I have the soft parts of three young specimens, of
which the sex is not determined, and five males, and four barren females.

Color of the distal part of the foot dark gray, this color sharply marked off
from the whitish basal part. The rest of the soft parts whitish.

Anal o])ening closed above; the closed part about four times as long as the
open, which is slit-like, shorter (about three-fourths) than the branchial opening,
and sejiarated from the latter by a solid mantle-connection. Branchial opening
short, with small, but distinct, papillae. Palpi rather large, subtriangular, lower
margins strongly convex, posterior margins with a short connection at base.

ortmann: south American naiades.

541

Gills long and wide, posteriorly of about the same width, but anteriorly the
inner is wider. Outer gill with margin curved, much narrower anteriorly, its
anterior end near the highest point of the mantle-attachment-line. Margin of
inner gill nearly straight in the middle, anteriorly this gill is a little narrowed,
its anterior end immediately behind the palpi. Inner lamina of inner gill entirely
connected with abdominal sac.

Non-marsupial gills with scattered, short, interrupted interlaminar connec-
tions, elongated in the direction of the gill-filaments. The 7narsupium of the
female (pi. XLVII, fig. 1) is in the inner gill, leaving free a small anterior and
posterior section, about one-fourth in front, and a little more behind, so that the
marsupium is located in the middle of the gill, but slightly more anteriorly. The
interlaminar connections (See also pi. XLVIII, fig. 4) are developed as rather
regularly interrupted septa, forming intercommunicating, incomplete water-tubes.
In the middle and at the base of the marsupium, the connections fall also into
irregular transverse lines, with a suggestion of quincuncial arrangement. In
young females, the marsupial part of the inner gill is considerably smaller.

It should be noted that the non-marsupial gills in this species are more nearly
normal than in D. gratus.

27. Diplodon rotundus Spix (1827).

Diplodoii rotu7idum and Utiio rotundus Spix & Wagner, 1827, p. 34, PL 25, figs. 3, 4.
Unio rotundus Sowerby, XVI, 1868, PI. 72, fig. 369; Von Ihering, 1890, p. 169, PI.
9, fig. 10; 1910, p. 139.

Diplodon {Cydomya) rotundus Simpson, 1900, p. 886; 1914, p. 1282.

Type-locality . — Southern Brazil.

Other Localities. — Rivers of eastern Brazil (Von Ihering, 1890); Rio S. Fran-
cisco, Villa Nova, Sergipe, Brazil (Von Ihering, 1910); Rio Paraguassu, Bahia,
Brazil (Von Ihering, 1893, p. 115; 1910); Rio Parahyba do Sul, Rio de Janeiro,
Brazil (Von Ihering, 1893, p. 115).

New Locality. — Rio S. Francisco, Bom Jesus da Lapa, Bahia, Brazil (J. D.
Haseman coll., December 17, 1907). Two odd right valves.

In the Carnegie Museum there is another, coinjilete specimen from the Hart-
man collection without locality.

Distribution. — Known from the lower and middle part of the Rio S. Francisco,
Rio Paraguassu, and Rio Parahyba do Sul in eastern Brazil.

According to the descriptions given by Von Ihering and Simpson my specimens
belong here, but my material is too scanty to give a full account of the species.

542

MEMOIRS OF THE CARNEGIE MUSEUM.

However, it should be said that of all species of the group this one most nearly ap-
])roaches the circular outline. It has been placed by Simpson, for this reason, in
the subgenus Cyclormja, but, as has been indicated already by Von Ihering, there
are siiecimens with the anterior part of the ventral margin less convex. The
figures of the type (Sjiix and Von Ihering) clearly show the oblique shape of the
sliell, with the highest part more posterior, at, or behind, the posterior end of the
ligament; and there is thus no question that this species falls into the same group
with fontamianus and grains.

Von Ihering (1893, p. 93) emphasizes the separation of the anterior protractor-
scar from the adductor-scar. In my specimens, this separation is not so distinct,
but only partial, so that we must not lay too much stress upon this character.

Measurements.

Loc.

Length.

Heiglit.

Depth.

Beaks.

Greatest Height.

Bom
.Jesus .

44 mm.

38 mm. =86 pr. ct . of L.

20 mm. =45 pr. ct. of L.

at 1 1 mm. =25 pr. ct. of L.

at 27 mm. =61 pr. ct. of L.

V

.55 “

48 “ =87

22.5 “ =41

12.5 “ =2.3

34 “ =62

Bom
.Jesus .

06 “

57 “ =86 “

32 “ =48

15 “ =23

41 “ =62

(V.lher.

1800)..

38 “

.32 “ =84

16 “ =42

10 “ =26

According to Von Ihering (1910, p. 139), this species reaches the length of
75 mm.

Subgenus; Cyclomya Simpson (1900).

As has been stated, I restrict this subgenus to those shells which have a sub-
rotund outline, with the greatest height located at about the middle of the shell,
that is to say, vertically below the middle of the ligament. The posterior part
of the shell is thus rather short, the shell is not obliciuely subtrapezoidal, as in the
species of the elli/pticus-gYowyi , but rather pentagonal in outline. The species at
hand is rather large and heavy, and belongs to the largest forms known in the genus.

28. Diplodon (Cyclomya) paranensis (Lea) (1834).

Shells: Plate XXXIX, figs. G, 7.

IJnio paranensis Lea, Obs. L, 1834, PI. 14, fig. 42; D’Orbigny, 1843, p. 603;

SowERBY, XVI, 18GG, PI. 51, fig. 268; Corsi, 1901, p. 451.

Diplodon {Cyclomya) ptaranensis Simpson, 1900, p. 887; 1914, p. 1284.

Unto rweturnus Lea, Obs. X, 1863, PI. 42, fig. 288.

Diplodon {Cyclomya) noclurnus Simpson, 1914, p. 1285.

Unio Paraguay anus Von Martens, 1895, p. 34.

Type-locality. — Rio Parana.

ortmann: south American naiades.

543

Other Localities. — Rio Uruguay (D’Orbigny) (Lea, nocturnus); Rio Parana up
to above Corrientes (D’Orbigny); Rio Paraguay (Von Ihering, 1893, p. 119);
Rio Paraguay, 25° S. Lat. (Von Martens) (This is near Asuncion, Paraguay).

New Localities. — Rio Paraguay, Corumba, Matto Grosso, Brazil (H. H.
Smith coll.). One large complete shell, one small left valve. In swamp of Lam-
bare, five miles below Asuncion, Paraguay (J. D. Haseman coll., March 31, 1909).
One right, one left valve. Rio de la Plata, San Isidro, Argentina (20 km. North of
Buenos Aires) (A. Windhausen coll., Januarj^ 1917). Five specimens with soft
parts, males and barren females.

Distribution. — Rio de la Plata, Parana, and Paraguay, from near Buenos
Aires up to Matto Grosso, Brazil; Rio Uruguajc

Description of Shell. — Shell solid and rather thick, large (length up to and
over 100 mm.). Outline angularly rounded, or broadly ovate, little oblique.
Height generally over 80 pr. ct. of the length, rarely less (lowest figure known 76
pr. ct., but this stands rather isolated, being found in a specimen measured by Simp-
son). Valves closed, or very little gaping in front. Dorsal margin gently curved
or straight (when young), descending anteriorly and passing into the anterior
margin, without forming a distinct angle. Posteriorly the dorsal margin forms
a more or less distinctly rounded angle with the posterior margin. The latter
descends obliquely and is straight, or even a little concave; then it curves around
in a sharp curve into the postero-ventral margin, this curve forming a blunt posterior
point of the shell, well elevated above the basal line. The ventral margin has a
sharp curve near its middle, forming a blunt, projecting angle at, or a little behind,
the middle of the shell. The highest part of the shell is at 50 to 60 pr. ct. of the
length. This ventral angle is generally distinct, but may be indistinct. From
this projection the lower margin slopes upward in either direction, backward and
forward, but the anterior part ascends more strongly, and is longer than the pos-
terior, and is often almost straight part of the way, before it finally curves up into
the anterior margin. The shell thus appears somewhat narrower anteriorly than
posteriorly, and has a rounded pentagonal shape, the five angles being formed by:
1, the beaks; 2, the upper posterior angle; 3, the posterior end; 4, the middle of the
lower margin; 5, the anterior end.

Valves not much swollen, moderately convex over the disk, but slightly flat-
tened on both sides of a submedian more convex ridge. The latter runs toward
the projection of the lower margin, but is very faintly marked. Posterior ridge
rather distinct on account of a distinct radial depression behind it. This depres-
sion makes the posterior slope much compressed, like a narrow wing; The slight

544

MEMOIRS OF THE CARNEGIE MUSEUM.

emargination of the posterior -margin corresponds to this depression. Diameter
40 to 46 pr. ct. of the length (according to Lea’s figure a little less, about 38 pr. ct.)
Beaks not swollen, and very slightly prominent beyond the hinge-line. Beak-
sculpture variable, but always more or less heavy, covering from 15 to 27 mm. of
the shell. There are about a dozen or more radial bars in front of the posterior
ridge, of which the anterior and jiosterior ones are narrow, while those in the
middle are thick and blunt. These ridges are very irregular; some anastomose,
smaller ones may be intercalated between the larger ones, or they may be connected
laterally. In some specimens the beak-sculpture is much less developed, but
traces of the heavy ridges are always visible. Toward the lower margin, the beak-
sculpture, when well developed, stops suddenh". In addition, there are upon the
jiosterior slojie a number of fine, oblique wrinkles, but these are only well-developed
in young individuals. In the larger specimens there is a short lunula, which is
narrow or very narrow.

Epidermis smooth, but with numerous irregular concentric wrinkles, which are
always crossed by more or less distinct radially impressed lines, forming here
and there low ridges. Where the beak-sculpture is well preserved, it is seen that
this radial sculpture is not a direct continuation of the beak-sculpture, but is in-
de])endent of it, beginning before the latter ends. In fact, the beginning of this
sculpture causes, in part, the irregularities of the beak-sculpture. Color of epi-
dermis yellowish brown to dark brown, generally darker posteriorly, without any
distinct color markings. Sometimes there are greenish olive tints towards the
beaks.

Hinge-line gently curved. Idgamental sinus shallow, over the middle of the
lateral teeth. Lateral teeth strong, moderately long, one in right, two in left
valve, descending jiosteriorly. In the youngest specimens, the upper margin is
somewhat elevated above the posterior ends of the laterals, while in the others
it is less elevated, almost parallel to them. Pseudocardinals strong, ragged, com-
pressed, running obliquely forward and downward, straight or curved. In the
right valve there are two iiseudocardinals, the posterior higher and stronger,
much cut up. In the left valve there is one strong tooth, much cut up, and some-
times a much smaller one behind it, which, however, may be absent. There is
much variation in the raggedness of the pseudocardinals.

Cavity of beaks and shell rather shallow. Nacre whitish or lurid, iridescent.
Anterior adductor-scar well impressed, subcircular, or subelliptical. Anterior
retractor-scar small, rounded, above the adductor-scar and separated from it.
Anterior iirdtractor-scar rounded, connected with adductor-scar. Posterior ad-

ortmann: south American naiades.

545

ductor-scar faintly impressed, almost pear-shaped, subovate or siibtriangular,
with an upper triangular process formed by the posterior retractor-scar. Mantle-
line distinct, remote from the margin about one-fourth to one-fifth (or less) of the
height of the shell. Dorsal scars about five (more or less), in beak-cavity in an
irregular, longitudinal row, sometimes shifted a little toward the narrow hinge-

plate.

Measurements.^!

No.

Sex.

Length.

Height.

Diameter.

Beaks at

Greatest Height at

Fig.

% L.

% L.

% L.

% L.

PI.

Corumba . . .

45 mm.

36

mm. = 80

20 mm. = 44

16

mm.

= 36

23

mm. = 51

XXXIX

S. Isidro ....

9

76

U

66

“ =87

33 “ -43

20

H

= 26

39

“ =51

f. 7.

Asuncion . . .

85

u

69

“ =81

34 “ =40

25

U

= 29

47

“ =55

PI.

S. Isidro a. . .

9

86.5

a

74.5

“ =86

36 “ =42

23

li

= 27

45

“ =52

XXXIX

Do. h. . .

9

90

i i

77

“ =86

40.5 “ =45

25

n

= 28

45

“ =50

f. 6.

Do. c. . .

91

u

77

“ =85

42 “ =46

27

i (

= 30

47

“ =52

Do. d. . .

92

u

80

“ =87

39.5 “ =43

27

u

= 29

55

“ =60

Corumba . . .

96

u

81

“ =84

41 “ =43

24

u

= 25

50

“ =52

Lea’s figure .

86

u

77

“ =90

33 “ =38

29

({

= 34

46

“ =53

Simpson ....

92

u

78

“ =85

38 “ =41

Do

100

ii

76

“ =76

47 “ =47

D’Orbigny. .

101

n

= 86

-37

Remarks. — Lea has described two other large, subrotund species from the
Uruguay River: nocturnus and funebralis (Obs. X, 1863, PI. 42, fig. 288, and PI. 41,
fig. 286). Of these, funebralis is similar in general shape and proportions, with
the exception of the much greater compression of the valves. The diameter is.
only 30 pr. ct. according to the figure and the measurements given by Simpson
(1914 p. 1284), which falls far below of any of the measurements known for paranen-
sis. In addition, in funebralis, the mantle-line is unusually far remote from the
margin of the shell, and, according to our present knowledge, we must consider
these characters as sufficient to separate the two species.

U. nocturnus was united with paranensis by Simpson in 1900, but in 1914
he separated them again. The proportions of nocturnus are:

Length.

Height.

Diameter.

Beaks.

Greatest Height.

Lea’s
figure . . .

72 mm.

60 mm. =83 pr. ct. of L.

30 mm. =41 iir. ct. of L.

at 21 mm. =29 pr. ct. of L.

at 37 mm. =51 pr. ct. of L.

Simpson’s
measur. .

72 “

60 “ =83

28 “ =30

Thus there is no essential difference from paranensis, and according to Simp-
son’s account, I can find only a difference in the color of the epidermis, which is

More care must be taken in tliis species than in others to place the ligament horizontally, ami to
measure parallel and vertical to it. The greatest height of the shell is hard to locate, and allowance
should be made for this.

546

MEMOIRS OF THE CARNEGIE MUSEUM.

said to be bottle-green, almost black posteriorly, in nocturnus. I do not think that
this is sufficient to separate nocturnus from paranensis, since in the latter olive-
green tints may also be noticed. This is seen chiefly in a specimen from the Hart-
man collection, without locality, preserved in the Carnegie Museum. Moreover,
the original description of Lea’s paranensis mentions this color.

The variability of the beak-sculpture of this species is remarkable. It has
never been described in detail, except that it consists of rather heavy radial bars.
In most of my specimens these bars are rather short (hardly more than 15 mm.
long), but in three (all isolated valves, two from Asuncion, one young from Corum-
ba) it extends farther, 25 to 27 mm., and chiefly in the young one (PL XXXIX,
f. 7) this is very striking, since in this case the beak-sculpture extends over nearl}^
half of the shell. I see, however, no other difference in these specimens, and even
among them the bars are not uniform in length. I believe that U. paraguayanus
Von Martens is founded upon such specimens. The dimensions of this fall easily
within the range of variation of paranensis. According to Von Martens they are:
length 102 mm.; height 82 mm. = 80 pr. ct. of length; diameter 47 mm. = 46per. ct.
of length. The location of the beaks is given as three-fourths of the length,” which
probably corresponds to 25 pr. ct. of length, we measuring from the anterior ex-
tremity, the author having reversed the jirocedure.

Von Martens compares his species with nocturnus, but says that it is larger,
with stronger sculpture (meaning ajiparently the beak-sculpture), and larger
hinge-teeth, but all these characters are unreliable, and do not distinguish it from
paranensis.

Anatomy. — The soft parts of two males and three barren females from San
Isidro are at hand.

Color whitish, distal part of foot gray.

Anal opening closed above ; the closed part being two to three times as long as
the open, which is slit-like, and shorter than the branchial opening. Anal and
branchial separated by a solid mantle-connection. Branchial opening with small,
but distinct, papillae. Paljii subtriangular, lower margins curved, posterior margins
connected for about one-third of their length.

Gills moderately wide, posteriorly of about the same width, but anteriorly
the inner gill is much wider. The shape of the latter is subtrapezoidal, that of
the outer gill subtriangular. Anterior end. of outer gill near the highest point of
the mantle-attachment-line, that of the inner gill immediately behind the palpi.
Inner lamina of inner gill entirely connected with abdominal sac.

Structure of non-marsupial gills as usual, with few and scattered interlaminar

ortmann: south American naiades.

547

connections. Marsupiu7n of the female located in the inner gill, but not occupying
all of it, leaving about one-fourth or a little less free at the anterior and the posterior
end. In younger specimens the marsupial part is smaller, but also has a median
position. Interlaminar connections of the marsupium forming interruiited septa,
without any distinct transverse or quincuncial arrangement.

Final Remarks on the Genus Diplodon.

It is not claimed that the above arrangement of the species of Diplodon should
be regarded as in an,y sense final. Even the two subgenera, Diplodon and Cydoniya,
cannot be sharply separated: they are intimately connected with each other,
and Cydomya is allied with the ellipticus-gi'oup of Diplodon through species like
rotundus, fontainianus, and gratus. The other groups cannot be very sharply
defined according to shell-characters. It is to be hoped that the anatom}^ may
furnish better criteria for the grouping of the species; but unfortunately not enough
species are known from this iioint of view, and some of those known are not known
fully enough. The characters of the glochidia need further special study. The
following facts, however, may be emphasized :

1. In six species of the diilensis-groiip {frenzeli, imitator, simillinius, vicarius,
decipiens, paidista), the marsupium has a distinct tendency to move forward in
the inner gill, and in three of these {simillimus, vicarius, paidista), it is entirely
anterior to the middle of the gill. In five of these species, the structure of the
interlaminar connections is interrupted seiitiform, or jiartly reticulate, but in one
species {decipiens) continuous septa are formed. The glochidia of all these sjiecies
have hooks.

The tendency of the marsupium to move forward is found in addition only in
two species of the ellipticus-gvoup (gratus and deceptus).

2. In the diarruanus-gmwp (charruanus, piceus, uruguayensis, hildcc), the
marsupium is rather large and lies in the middle of the gill; sometimes (hildw) it
is smaller; and in two cases {charruanus and hildai) it has a tendency to move back-
ward. In the lacteolus-gm\vp (two species, burroughianus and niogymirini) , the
marsupium is also located slightly more posteriorly. In most of these species,
the interlaminar connections form interrupted septa, in part reticulated. But
there is one exception, for mogyniirirn has continuous septa. This structure is
thus only known in two species {niogyniirvm and decipiens) belonging to different
groups.

3. In the ellipticus-growp , the marsupium may be in the middle of the gill
(enno), slightly posterior (herthcc), or slightly anterior {gratus, deceptus). Its

548

MEMOIRS OF THE CARNEGIE MUSEUM.

structure may be prevailingly reticulate {heriim, enno), or prevailingly septiform
{gratus, deceptiis).

4. D. paranensis has the marsupium in the middle half of the gill, and in-
terrupted septiform structure is present.

5. In eight species, hooks are known to be present on the glochidia. Of
these six belong to the c/u7cn.sfs-group, one to the charruoMus-group (piceus), one
to the lacteolus-gi'oiip {niogymirim). Glochidia without hooks (but possiblj'^ im-
mature) were found in charnumus and herthce {charruanus- and ellipticus-grovip^) .
Two species have margined glochidia, hasemani of the hylceus-group, and hildce
of the charruanus-group.

For the present, these facts do not furnish any clue as to the relationship of
the species or groups, they even are rather confusing, partly uiisetting the divisions
arrived at by the study of the shell. But they should be carefully recorded,
because additional material may throw more light on the problem.

Genus CASTALINA Von Ihering (1891).

Von Ihering, 1891, p. 478; 1893, p. 73; Simpson, 1900, p. 865; 1914, p. 1204.

Type-species. — C. 7nartensi Von Ihering (designated by Simpson).

The chief characters of this genus are found in the general shape of the shell,
which is subfriangular or subquadrate, with a well developed posterior ridge and
a subtruncated posterior slo]ie, which, however, is somewhat elevated in the middle.
In addition, the beaks are rather elevated, the interdentum is well developed, form-
ing a rather deep beak-cavity. The hinge-teeth often are ])rovided with parallel
ridges.

The anatomy is similar to that of Diplodon. However, there is a tendency to
close the liranchial oiiening in front, yet this is not always the case, so that this
character is variable, not only sjiecificalljq but also individually.

The genus has been well treated by Von Ihering (1893), and a key for the
species has been given (p. 83). Simpson (1914 p. 1205) also gives a key, but C.
undosa should be excluded; it is a CasUdia.

29. Castalina nehringi Von Ihering (1893).^^

Diagram of soft parts: text-fig. 2, p. 456; Anatomy of gills: PI. XLVII, fig. 2; Section
of gills: Plate XLVIII, fig. 5; Glochidimn: text-fig. 41, p. 469.

Type-loccdity. — Rio Piracicaba, Sao Paulo, Brazil.

Of both C. nehringi and niartcnsi the specific names were first mentioned by Von Ihering in 1891
(p. 477), blit as nomina nnda without descriptions.

ortmann: south American naiades.

549

Material Represented in the Carnegie Museum. — Rio Piracicaba, Sao Paulo,
Brazil (Von Ihering, donor, cotypes or topotypes). Two specimens. Rio Tiete,
Salto das Cruzes, Sao Paulo, Brazil (J. D. Haseman coll., September 22, 1908).
Four specimens, two of them with soft parts. Rio Tiete, 25 miles above Itapura,
Sao Paulo, Brazil (J. D. Haseman coll., September 27, 1908). Eight specimens,
six of them with soft parts.

Distribution. — Rio Tiete and Rio Piracicaba in Sao Paulo, headwaters of Rio
Parana.

Von Ihering has given a detailed description of this species, pointing out its
differences from the allied species. Pie believes that specimens with the beaks
more distant from the anterior margin are males. I cannot control this, since I
have no males among those of my specimens with soft parts, but the percentage
given for the location of the beaks in the males (28 to 30 pr. ct.) is not represented
in my measurements, and this would tend to confirm Von Ihering’s observation.
It is to be noted that my smallest specimens (Nos. 1 to 5 from Itapura) exceed in
height the figures given by Von Ihering, and thus young shells are proportionally
higher: the older ones become longer on account of a prolongation of the posterior
end of the shell (best seen in No. 6 from Itapura).

Measurements.

No. Sex.

Length.

Height.

Diameter.

Beaks.

Itapura

1

9

51

mm.

46 mm. =90 lu-.

ct. of L.

26

mm. =51 pr. c

h. of L.

at 9

mm. =18 pr.

ct. of L.

Do

5

9

54

40.5 “ =80

29.5

“ =55

11

“ =20

Do

.3

9

55

48 “ =87

26

“ =47

10.5

“ =19

Salto (las Cruzes

1

9

01.5

49.,5 “ =S0

31

“ =50

“

12

“ =20

Do

2

9

69

,55.5 “ =80

34

“ =49

15

“ =22

Itapura

0

9

81

04 “ =79

40

“ =49

15.5

“ =19

Anatomy. — I have the soft parts of eight specimens, all of which are females,
three of them gravid. One of the latter had eggs; one had immature, and the
third mature glochidia. For the breeding season the dates of collection (Sejitember
22 and 27) should be noted. (Mature glochidia were found on Sept. 22.)

Von Ihering (1893, \). 79) describes the soft ]iarts of two males. Of these,
one had the branchial opening closed in front, the other open. This is the species
of Castalina, to which I have referred previously (Ortmann, 1911, p. 118). In
none of my females is there an anterior mantle-connection in front of the branchial
opening. Therefore this seems to be the normal condition in this species, although
it should be born in mind that this connection at its best is very slight, and might
be easily torn by rough handling.

The anal opening is described by Von Ihering as also differing in his two

550

MEMOIRS OF THE CARNEGIE MUSEUM.

specimens. He mentions the presence in one of a siipra-anal opening; but is not
quite ])ositive that this is natural. According to my material, the anal is always
closed above, without forming a supra-anal (text-fig. 2, s, p. 45G). The soft parts
have the following characters:

Anal opening closed above; open part (text-fig. 2, a p. 456) short, somewhat
shorter than the liranchial, slit-like, its inner margin indistinctly crenulated or
smooth. Closed ])art (s) about three to four times as long as the open part. A
supra-anal canal extends all the way under the closed part (above the rectum),
ending blindly aliove. Mantle-connection between anal and branchial openings
well developed (text-fig. 2, t). Branchial ojiening (6) with well developed papillse
on inner margin, which show some irregularities at the anterior end of the opening,
but in none of my specimens are distinct traces of a connection of the mantle margins
in this region visible. Paliii (/i) large, subtriangular, almost falciform, with long
and curved lower margins, and short posterior margins, the latter connected for
about one half of their length.

Gills (text-fig. 2, i, o) moderately wide, the inner (t) distinctly wider, chiefly
anteriorly. Outer gill (o) with gently curved edge, its anterior end near the highest
point of the mantle-attachment-line. Inner gill with the edge almost straight,
anteriorly a little narrower, and broadly attached, the attachment occupying all
of the space between the anterior end of the outer gill and the palpi (/i). Inner
lamina of inner gill entirely connected with abdominal sac. In the female (PI.
XLVII, fig. 2) the inner gill is marsupial, but only a section of the gill possesses this
character, with the interlaminar connections distinctly arranged in interrupted
septa (See also Pl.XLVIII, fig. 5). The marsupial part is rather small in younger
specimens (such as the one figured on PI. XLVII, fig. 2), lying immediately behind
the middle of the gill; but it is larger in older siiecimens, lying practically in the
middle, leaving free aliout one-third anteriorly as well as posteriorly (text-fig. 2, i).
The non-marsupial gills have remote, incomplete, interrupted septa, and the septi-
form structure is more evident than it generally is in Diplodon (See outer gill,
PL XLVII, fig. 2).

When charged, the eggs or glochidia do not form placenta-like masses. The
fully develojied glochidium (text-fig. 4, 1, p. 469) is subtriangular, with a lower point
situated about in the middle of the lower margin, and with distinct hooks at this
point, which differ from those seen in certain species of Diplodon in that they are
shorter, and broader at the base. L. 0.26 mm.; H. 0.24 mm.; hooks: 0.06 mm.
Thus the glochidium is rather small.

ortmann: south American naiades.

551

30. Castalina martensi Von Ihering (1893).

Von Ihering, 1893, p. 81, PI. 3, fig. 5; Simpson, 1900, p. 865; 1914, p. 1205.

Type-locality . — Rio Camaquam, Rio Grande do Sul, Brazil.

New locality. — Rio Jaciiliy, Cachoeira, Rio Grande do Sul, Brazil (J. D. Hase-
man coll., January 27, 1909). Three complete shells and one right valve.

Distribution. — Our new locality extends the range of this species northward.
The Rio Jacuhy belongs to the Guahyba-system, but both the Guahyba and Cama-
quam flow into the Lagoa dos Patos. Von Ihering says that he did not find this
species north of the Rio Camaquam.

This species has been sufficiently well described by Von Ihering. Our youngest
specimen shows well developed beak-sculpture, corresponding to the description.
The variations mentioned by Von Ihering in the crenulation of the lateral teeth
are also seen in our specimens.

Measurements.

Length.

Height.

Diameter.

Beaks.

26 min.

21.5 mm. = 83 pr. ct. of L.

13.5 mm. = 52 pr. ct. of L.

at 8.5 mm. = 33 pr. ct. of L.

48 “

38 “ =79

24 “ =50

12 “ =25

48 “

40.5 “ =84

26 “ =54

13 “ =27

75 “

55 “ =73

34 “ =45

21 “ =29 “ Largest speci-

men of Von Ihering)

The height ranges according to Von Ihering from 70 to 79 pr. ct., and the diam-
eter from 45 to 50 pr. ct. The location of the beaks is from 21 to 29 pr. ct. Thus
our specimens are comparatively higher and more swollen. As far as concerns the
height, this is certainly due to the smaller size of my shells, since a similar change
in the proportional height with age has been observed in C. nchringi.

Anatomy. — Of this species Von Ihering (1891, }). 477) says that in most cases
(seven out of eight) the branchial opening is closed in front. I do not possess the
soft parts.

31. Castalina psammoica (D’Orbigny) (1835).

Unio psammoica D’Orbigny, 1843, p. 608, PI. 71, figs. 4-7; Sowerby, XVI, 1868,

PL 93, fig. 507.

Castalina psammoica Von Ihering, 1893, p. 79; Von Martens, 1894, p. 164;

Simpson, 1900, p. 866; 1914, p. 1206; Haas, 1916, pp. 9, 47.

Type-locality. — Rio Parana, Itaty, above C’orrientes, Argentina.

Other Localities. — Province Santa Fe, Argentina (D’Orbigny) (farther down the
Parana); Rio Paraguay, near mouth of Rio Apa (Von Ihering) (in Paraguay, at
Brazilian boundary); Paraguay (Von Martens); Rio Uruguay, Salto Oriental,
Uruguay (Haas).

552

MEMOIRS OF THE CARNEGIE MUSEUM.

New Localities. — Rio Uruguay (in mud), Uruguayana, Rio Grande ^ do Sul,
Brazil (J. D. Haseinan coll., February 5, 1909). One young male with soft
parts. Rio de la Plata, San Isidro, 20 km. north of Buenos Aires, Argentina (A.
Windhausen coll., January 1917). One male with soft parts.

Distribution. — From the La Plata near Buenos Aires up to the Rio Parana and
Rio Paraguay in Paraguay, and also in the Rio Uruguay.

The descriptions given by D’Orbigii}^, Von Ihering, and Simpson agree well
with our specimens, but it should be mentioned that the beak-sculpture sometimes
covers less than half of the adult shell. The hinge-teeth are very variable, but
Von Ihering’s account represents the normal condition. The laterals are said
to be smooth or finely smoothly striated. In my young specimen they have dis-
tinct, vertical, irregular, and granular ridges; in my larger specimen they are ir-
regularly granular and crenulated, but have no distinct vertical ridges.

According to Von Ihering, the maximum length is 70 mm.; according to D’Or-
bigny, 75 mm. The height according to the former is 77 to 80 pr. ct.; according to
the latter, 76 pr. ct.; the diameter is 50 and 49 pr. ct. respectively. The beaks are
located, according to Von Ihering at 17 to 23 pr. ct. of the length.

Measurements.

Localities.

Sex.

Length.

Height.

Diameter.

Beaks.

Uruguayana

c?

44 mm.

35 mm. =80 pr. ct. of L.

23 mm. =52 pr. ct. of L.

at 10.5 mm. =24 pr. ct. of L.

San Isidro

cf

62 “

50 “ =81 . “

37.5 “ =60

14 “ =23

Thus our larger specimen is unusually swollen.

Remarks. — This species somewhat resembles the genus Castalia, chiefly on
account of the strong development of the beak-sculpture. However, according to
the posterior slope, which is distinctly elevated along the upper posterior margin,
and also according to the structure of the lateral teeth, which have the vertical
ridges poorly, or not at all, developed, it is a Castalina. ' In Castalia furthermore the
diameter of the shell generally is much greater than the maximum (60 pr. ct.) ob-
served in the present species.

Anatomy. — Soft parts of two males at hand. Color whitish.

Anal opening slit-like, closed above; closed part three to four times as long as
the open, the latter shorter than the branchial opening, and separated from it by
a solid mantle-connection. Branchial with small papillae. In the smaller specimen,
the branchial opening is closed in front by a firm union of the mantle-margins,
but in the larger specimen there is no such connection, and thus in this species
this character is variable. Palpi rather large, nearly subfalciform, with the lower

ortmann: south American naiades.

553

margins long and strongly curved, posterior margins short, and united at base for
about one-fourth of their length.

Inner gills much wider than the outer, chiefly in front; only for a short
distance behind have they an equal width. Outer gill with curved margin, anterior
end at highest point of mantle-attachment-line. Inner gill with margin nearly
straight, broadly attached anteriorly, and anterior end close to the palpi. Inner
lamina of inner gill entirely connected with abdominal sac. Structure of non-
marsupial gills normal, with scattered interlaminar connections, assuming here and
there the shape of short, weak septa.

Genus CASTALIA Lamarck (1819).

Castalia Lamarck, 1819, p. 66; Pilsbry, 1911, p. 610.

Tetraplodon Spix, 1827, PI. 25; Pilsbry, 1893, p. 90; Simpson, 1900, p. 863; 1914,

p. 1194.

Pilsbry (1911) has brought to light the fact that Savigny’s genus Castalia
{Vermes) is not to be dated from 1817, but that it is later (certainly not before 1820,
probably as late as 1826), so that Castalia applied by Lamarck (1819) to the present
genus has to stand.

Castalia is distinguished by the triangular outline, with high beaks, well de-
veloped posterior ridge, great obesity of the shell, truncated posterior slope, and
well developed beak-sculpture, which extends over a large part of the shell. It
is to be noted, however, that the truncation of the posterior slope often varies with
age, so that in very young specimens it is somewhat elevated in the middle, giving
an outline to the shell approaching the subtrapezoidal or subrhomboidal. With
one exception the species resemble each other very closely. This exception is
C. undosa, which differs from all the rest in having strong, oblique folds, or ribs, upon
the posterior slope.

My material of Castalia is rather insufficient, and I have had great difficulties
in determining the species. The key given by Simpson (1914) is useless for the
reason that the type-species {amhigua) has been misunderstood. The latter has
been redescribed and figured by Von Ihering (1910),^^ which fact had been neglected
by Simpson. At the same time Von Ihering has given an elaborate key for the
species. However, this also is unsatisfactory, and 1 am afraid that Von Ihering
has not paid proper attention to the changes undergone by the shell with advancing

In this connection it should be mentioned that Wyatt, T. (A Manual of Conchology, 1838, p. 65,
PI. 11, fig. 5) has given a recognizable figure of C. anibigua, which undoubtedly represents the genuine
amhigua, as determined by Von Ihering).

554

MEMOIRS OF THE CARNEGIE MUSEUM.

age. Ill trying to use this key, I generally met with the difficulty that my specimens
were intermediate between the alternatives given in it. But, as I said, my material
is too scanty to firing order out of this confusion, yet I shall try my best to give an
account of the material at hand.

The geographical distribution of the species of this genus is interesting. Ac-
cording to Von Ihering (1910, p. 130) their metropolis is in the Amazon-drainage,
whence the forms extend into Guyana and the upper Parana-drainage. But on
account of the great uncertaint}^ iirevailing' with regard to the various species no
details can be given.

32. Castalia acuticosta Hupe (1857).

Glochidium: Text-fig. 4, m, p. 469.

Castalia acuticosta Hupe, 1857, p. 77, PI. 14, fig. 3; Sowerby, XVII. 1869, PI. 3,

fig. 12.

Telraplodon acuticosta AVin Ihering, 1910, p. 128.

Type-loccdity. — Brazil. (Von Ihering says that Castelnau collected this
species in Goyaz, but I cannot find this fact mentioned in Hupe’s paper.)

Other Localities. — Rio Araguay, Goyaz, Brazil (Von Ihering, 1910, p. 135) ;
Lagoa do Coral, Goyaz, Brazil (Von Ihering) (drainage of Rio Araguay, tributary
to Tocantins).

New Localities. — Sand-bar of Rio Tapajos, Santarem, Para, Brazil (J. D. Hase-
man coll., December 6-12, 1909). Eight specimens, one of them a male with soft
])arts. Center of Rio Guapore, near Rio Sao Simao, Matto Grosso, Brazil (J. D.
llaseman coll., July 20, 1909). Three siiecimens, with soft parts, male, and gravid
females. Rio Guapore, Sao Antonio de Guapore, Matto Grosso, Brazil (J. D.
llaseman coll., July 31, 1909). Four specimens. Rio Machupo, San Joaquim,
Bolivia (J. D. Haseman coll., September 1, 1909). Three 3mung specimens, with
soft parts, probably males.

In addition, the Carnegie Museum possesses a fine specimen from the Juny
collection, without locality.

Distribution. — Southern tributaries of the Amazon, from Tocantins and Ara-
guaj" in Go,yaz, and Rio Tapajos in Para, to the Madeira (Guapore and Machupo)
in Matto Grosso and Bolivia.

This species belongs to those with elevated beaks, although the beaks are here
far less prominent than in turgida, baro, and hanlcycma. It is a small species
(maximum length 35 rnni.); but that specimens about 25 mm. long are mature is
shown by the fact that at that size they may be gravid.

ortmann: south American naiades.

555

The outline is “subquadrate” according to Von Ihering. I should call it
rather subtrapezoidal or subrhomboidal, and this is due to the fact that the pos-
terior end is not much produced, and that the angle between the upper and the
posterior margins is somewhat prominent; consequently, the posterior slope is
not so much truncated as in other species, but a little elevated toward the upper
posterior angle. 0n account of the shortness of the shell, the beaks are com-
paratively remote from the anterior end.

Von Ihering has pointed out that there are variations in the beak-sculi)ture,
and I observe this also in my specimens. In most cases it is sharp,' the bars are
narrow and distinct, and extend to the lower margin of the shell. But sometimes
the bars are more rounded, less distinct, and may not reach the lower margin. A
few finer radial bars are generally present upon the posterior slope.

AIeasukements.

Localities.

No.

Sex.

Length.

Heiglit.

Diameter.

Beaks.

Santarem

2

?

18.5

nmi.

16

mm. =86 pr. ct. of L.

14

mm. =76 pr. ct. of L.

at 5.5

mm. =24 pr. ct. of L.

Do

3

20

18

“ =90

17

“ =85

0.5

“ =33

Guapore

2

9

24

20

“ =83

19

“ =79

6.5

“ =27

Do

1

9

24

20.5

“ =85

18

“ =75

6

“ =25

Do

3

c?

28

24.5

“ =88

21

“ =75

8

“ =29

Santarem

e

?

29

25

“ =86

22

“ =76

8.5

“ =29

?

?

30

27

“ =90

23

“ =77

10

“ =33

Hupe, text-figure

34

26

“ =76

23

“ =68

It should be remarked that Hupe’s figure of the diameter (half shell) does
not agree with his measurements in the text. This figure would give a diameter
of only 18 mm. ( = 53 pr. ct.,) while the text gives 68 pr. ct. Both height and diam-
eter of Hupe’s shell are less than in any of mine. Von Ihering gives the location of
the beaks at 39 to 41 pr. ct. of the length, which is much greater than the figures
of Hupe (31 pr. ct. according to the figure) and of my shells. But I believe that this
discrepancy is due to a different way of measuring: I always measure parallel to
the ligament.

Remarks. — My specimens certainly belong together; but I am not quite positive
that they might not be only the young stage of some other species {Jumlcyana or
haro). The fact that some of mine are gravid does not demonstrate that they are
adult or nearly adult, for we know that other Naiades become sexually niature,
when yet rather small. The further fact that those from Bantarem were found
associated with baro and hanleyana indicates that they may belong to these species.
However, I am unable to decide this question.

Anatomy. — Soft parts of two males and two gravid females (with glochidia) at
hand. The latter were found on July 20. Color of soft parts whitish.

556

MEMOIRS OF THE CARNEGIE MUSEUM.

Anal opening slit-like, closed above; closed part about four to five times as
long as the open, the latter sejiarated from the branchial opening by a solid mantle-
connection. Branchial opening with small papillie, closed in front by a mantle-
connection in the two males and in one of the females, while in the other female
this connection is missing (but it may be torn). Palpi moderately large, sub-
triangular, with curved lower margins, and the posterior margins united for about
one-fourth of their length.

Gills about equally" wide posteriorly, anteriorly the inner is the wider, of the
usual shape. ‘ Inner lamina of inner gill entirely connected with the abdominal
sac. Non-marsupial gills with scattered interlaminar connections. Marsupium
in the middle of the inner gill, leaving about one-fourth of the gill free at the anterior
end, and a little less at the posterior. Interlaminar connections interrupted, but
their arrangement cannot be clearly seen, since no barren females are at hand.

Glochidium (text-fig. 4, m, p. 469 ) subtriangular, with a point at about the
middle of the lower margin, and with hooks of the usual shape, but gather short.
L. and H. about 0.24 mm., hooks 0.05 mm. Thus the glochidium is rather small.

33. Castalia hanleyana Sowerby (1869).

Castalia hanleyana Sowerby, XVII, 1869, PL 1, f. 5.

Tetraplodon hanleyanus Von Ihering, 1910, p. 127.

Type-locality . — Unknown.

Other Localities. — Rio Araguaya, llha Bananal, Goyaz, Brazil (Von Ihering);
State of Para, Brazil (Von Ihering).

New Locality. — Sand bar of Rio Tapajos, Santarem, Para, Brazil (J. D. Hase-
man coll., December 6-12, 1909). Two odd valves (right and left).

Distribution. — It should be noted that this species is found, where also acuti-
costa occurs, in the Rio Araguaya and the Rio Tapajos.

My two valves agree well with the account given by Von Ihering, and with
Sowerby’s figure, but are smaller than the latter.

Measurements.

Lonstli.

Height.

1

Diameter.

1

Beaks.

42.. mm.

35 mm. = 82 pr. ct. of L.

28 mm. = 66 pr. ct. of L.

at 12

mm. = 28 pr. ct. of L.

36.5 “

31 “ =85

27 “ =74

10.5

“ =29

Remarks. — According to Von Ihering, the beaks are at 30 to 33 pr. ct., while my
specimens show lower figures. But this fact may be due to a different way of •
measuring. The shell is characterized by high beaks and its high, triangular (not

ortmann: south American naiades.

557

elongated) shape, which makes the beaks appear to be rather far back (in comparison
with baro). However, this may be only a more aged form of acuiicosta.

34. Castalia baro (Von Ihering) (1910)

Castalia ambigua Sowerby (not Lamarck), XVII, 1869, PI. 1, fig. 1.

Tetraplodon baro Von Ihering, 1910, p. 127; Simpson, 1914, p. 1198.

Type-locality. — Amazon River.

New Locality: Sand bar of Rio Tapajos, Santarem, Para, Brazil (J. D. Hase-
man coll., December 6-12, 1909). One complete shell, two odd valves, right and
left.

In addition the Carnegie Aluseum has two other fine specimens, one from the
Holland collection, labeled “River Amazon,” the other from the Jiiny collection,
without locality.

Distribution: Amazon and Tapajos Rivers.

This species stands very close to C. hmileyana, and differs only in its more
elongated shape, and, consequently, more anterior beaks. Von Ihering says that
the lower margin is straight (while it is convex in hanleyayia). But this feature is
somewhat variable, for it may be gently curved. However it is correct that in
baro the lower margin is more nearly straight than in hanleyana.

I should not be astonished if ha7ileyana and baro should finally prove to be
only the adult stages of acuiicosta. The fact that all three have been found by
Haseman associated at Santarem is in favor of this view.

About the dimensions of baro we only know that the beaks are at 16 pr. ct. of
the length. My specimens show a somewhat higher percentage, but not as high
as in hanleyana (where it is from 28 to 33 pr. ct.).

Measurements.

Localities.

Length.

Height.

Diameter.

Beaks.

Santarem . .

39 mm.

28 mm. = 73 pr. ct. of L.

27 mm. = 69 pr. ct. of L.

at 9 mm. =23 pr. ct. of L.

Do. . .

45.5 “

33 “ =73

30.5 “ =67

9 “ =20

Amazon . . .

48

35 “ =73

32 “ =67

10.5 “ =22

(.Juny) ....

52.5 “

39.5 “ =75

37 “ =72

10 “ =19

Santarem . .

55

CO

il

o

36 “ =65

10 “ =18

35. Castalia pectinata (Spix) (1827).

Tetraplodon pectinatum Spix, 1827, PI. 25, fig. 3; Von Ihering, 1910, p. 125.
Unio pectinatus Wagner, 1827, p. 32.

Castalia ambigua Von Ihering (not Lamarck), 1890, p. 162.

Type-locality. — Rio Sao Francisco, Minas Geraes, Brazil.

558

MEMOIRS OF THE CARNEGIE MUSEUM.

New Localities. — Amazon River, Para, Brazil (Hartman collection). Two
specimens. Sand bar of Rio Tapajos, Santarem, Para, Brazil (J. D. Haseman
coll., December 6-12, 1909). One odd (right) valve.

Distribution. — Known hitherto only from the Rio Sao Francisco, and only
from the description and figure of Sjiix and Wagner. Its discovery in the Amazon
and Tapajos in Para thus extends its range into the lower basin of the Amazon.

The characters of this species have been established by Von Ihering, and,
although it is hazardous to define an old, often misunderstood, and poorly figured
form, without having abundant material, the fact that our specimens well agree
with the account given by Von Ihering, demonstrates that he is right. The chief
characters of this species are found in the low beaks, the rather elongated outline,
and rather anterior position of the beaks. The beak-sculpture is described as
‘‘mostly” reaching the ventral margin. This is actually the case in one of our
specimens from the Amazon and in the half shell from the Tapajos. In the other
individual from the Amazon the radial bars are shorter and cover only a little
more than half of the shell.

IMeasurements.

Locality.

Letigtli.

Ileiglit.

Diameter.

Beaks.

Amazon. . .

28 mm.

21.5 mm. = 77 pr. ct. of L.

17 mm. = 61 pr. ct. of L.

at 6 mm. =21 pr. ct.of L.

Tapajos. . .

28 “

21 “ =75

18 “ =64

6.5 “ =23

Amazon . . .

31 “

23 “ =74

20 “ =65

8 “ =26i

Remarks. — Von Ihering gives the location of the beaks as 25 pr. ct.; my speci-
mens agree with this very well.

C.juruana (Von Ihering, 1910, p. 126) from the Rio Jurua (southern tributary
of the Amazon in western Brazil) is closely allied. It is founded upon a single
specimen, which differs chiefly in the location of the beaks (at 37 pr. ct.), and certain
other characters, which at least in part are connected with this. It may finally
prove to ‘be only a form of pectinata.

Our specimens have about the size given by Wagner in the text {1. c., p. 33)
28 to 31 mm.; while Spix’s figures (PI. 24, figs. 3, 4) are considerably larger (L.
over 40 mm.) . Thus they apparently are enlarged. C. juruana is larger, L. 46 mm.

36. Castalia inflata D’Orbigny. (1835).

Castalia inflata D’Orbigny, 1835, p. 43.

Castalia anibigua D’Orbigny, 1843, p. 598, PI. 72, figs. 4-6.

Castalia anibigua inflata Von Ihering, 1893, p. 88.

Castalia quadrilatera Von Ihering, 1893, p. 89.

ortmann: south American naiades.

559

Tetraplodon inflatus Von Ihering, 1910, p. 126; 1915, p, 12.

Type-locality. — Parana River, Corrientes, Argentina.

Other Localities. — Lower Parana River (Von Ihering) ; small tributaries of the
Parana (D’Orbigny); Rio Paraguay, near Rio Apa, Paraguay (Von Ihering); Rio
Paraguay, San Luis de Caceres, Matto Grosso, Brazil (Von Ihering).

Locality Represented in Carnegie Museum. — Rio Paraguay, San Luis de Caceres,
Matto Grosso, Brazil (J. D. Haseman coll.. May 24, 1909). Two specimens.

Distribution. — Positively known only from the Parana and Paraguay Rivers in
Argentina, Paragua}^, and Brazil. Specimens reported from the Amazon-drainage
in Bolivia probably do not belong here.

My identification of this sjiecies depends partly on Von Ihering’s key, partly
on D’Orbigny’s original description and figures of specimens from Corrientes,
and on the locality. It is a species with low beaks, the shell is oval, but for the
rest. Von Ihering’s key is insufficient. One of my specimens is rather small, and
the beak-sculpture reaches the lower margin; in the other, it falls short of it. The
beaks are located at 21 and 24 pr. ct. (21 pr. ct. given by Von Ihering). The shell
is about as long as in pectinata, and is thicker. The bars of the beak-sculpture
in my specimen are rather numerous, but not very sharp. The adductor-scars
agree with Von Ihering’s account. Thus I have no doubt that my specimen repre-
sents C. inflata, although the essential characters of this species are yet obscure.

Measurements.

Length.

Height.

Diameter.

Beaks.

D’Orbigny’s text

D’Orbigny’s fig. of specimen

29 mm.
36 “

65 “

23 mm. =79 pr. ct. of L.
26 “ =72

75

20 mm. =69 pr. ct. of L.
22.5 “ =63

60

at 6 mm. =21 pr. ct. of L.
8.5 “ =24

from Corrientes

35 “

25.5 “ =73

24 “ =70

8.5 “ =24

Von Ihering does not give detailed measurements, but only says that the maxi-
mum length is 51 (1893) and 60 mm. (1910).

37. Castalia undosa Von Martens (1885).

Shell: Plate XXXIX, figs. 8a-d.

Castalia undosa Von Martens, 1885, p. 148; Von Ihering, 1893, p. 84; Nehring,
1893, p. 165.

Castalina undosa Simpson, 1900, p. 866; 1914, p. 1207.

Tetraplodon undosus Ortmann, 1911a, p. 117, PI. 6, fig. 7, PI. 7, fig. 7 (anatomy).
Type-locality . — Rio Piracicaba, Piracicaba, Sao Paulo, Brazil.

560

MEMOIRS OF THE CARNEGIE MUSEUM,

Localities Represented in Carnegie Museum.. — Rio Piracicaba, Sao Paulo,
Brazil (Von Ihering donor, authentic specimens). One complete shell and one
odd (left) valve. Rio Tiete, silty river banks, twenty-five miles above Itapura,
Sao Paulo, Brazil (J. D. Haseman coll., September 27, 1908). Four specimens
with soft parts, one male and three females. Rio Tiete, Itapura, Sao Paulo, Brazil
(J. D. Haseman coll., September 28, 1908). Five specimens, one of them a male
with soft parts.

Distribution. — Rio Tiete and Piracicaba of Upper Parana-drainage in Sao
Paulo, Brazil.

This species has been well described by Von Martens and Von Ihering, and I
do not need to add anything, except that the dimensions vary considerably, the
height from 71 to 86 pr. ct. and the diameter from 53 to 66 pr. ct. of the length. On
the average our specimens are more swollen than the previous measurements would
indicate. The most characteristic feature of this species is the quite unique de-
velopment of the ribs of the posterior slope, which are much stronger than the
oblique ‘‘wrinkles” often found in species of Diplodon.

Sim]:)Son has put this species in the genus Casta.lina, but I do not see any
reason for this. The posterior slope undoubtedly has the characters of Castalia.
The species, indeed, stands isolated in the genus, and cannot be confounded with
the other species, and for this reason, probably. Von Ihering omitted it in his key
(1910).

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Figure.

?

42 min.

34 mni

= 81 pr. ct. of L.

20

mm. =62 pr. ct. of L.

at 10

mm. =24 pr.

ct. of L.

PI. XXXIX,

X

9

40.5 “

39

=84

27

“ =58

11.5

“ =25

fig. 8.

b

9

47

37.5 “

=80

31

“ =60

11.5

“ =24

c

9

51

38.5 “

= 75 “

31

“ =01

13

“ =25

a

o’

57

44

=77

33.5

“ =59

13.5

“ =24

O’

00

47

= 78

36

“ =00

14

“ =23

(Von

Martens) . .

50

48

= 80

30

“ =53

2/7

= 29

“

(Von Iliering)

00

50

= 70

36

“ =55

Anatomy. — The soft parts of two males and three barren females are at hand.

A descrijition has been previously given by myself (1911a, p. 117). However,
it should be added that in one female investigated the branchial opening is open
in front, liut probably torn. In another, a male, it is undoubtedly open; this is
an old individual, the largest at hand, and here also it may be that it has been in-
jured during life. In the other three specimens (one male and two females), the
branchial opening is closed in front. According to Von Ihering (1891, p. 476),
this opening is closed in 80 pr. ct. of the cases.

ortmann: south American naiades.

561

Anal opening closed above; closed part three to four times as long as the open
part, the latter slit-like, shorter than the branchial, and separated from the latter
by a solid mantle-connection. Branchial opening with papillae, closed in front
by a firm union of the inner mantle-edges, or, more rarely, open and without this
connection. Palpi rather large and slightly produced posteriorly, with strongly
curved lower margins. Their posterior margins are connected for about one-
third of their length.

Gills of the usual shape, the inner much wider in front than the outer, its
anterior end close to the palpi. Inner lamina of inner gill entirely connected
with abdominal sac. Non-marsupial gills with scattered interlaminar connec-
tions. Marsupium of the female located in the inner gill, but it does not occupy
all of this gill, but only a portion in the middle, leaving free a larger part anteriorly,
and a smaller posteriorly, so that the marsupium appears shifted slightly backwards.
Interlaminar connections of marsupium arranged in interru]3ted septa.

Genus Hyria Lamarck (1819).

Hyria Lamarck, 1819, p. 81; Simpson, 1900, p. 868; 1914, p. 1211.

This genus is characterized by the alate shape of the shell, possessing wings
both at the anterior and posterior end of the upper margin. In addition the
radial beak-sculpture is well developed, covering a considerable part of the shell,
the posterior ridge is rounded and broad, and the jiseudocardinals are much com-
pressed. The hinge-teeth may be somewhat corrugated, but they do not possess
distinct vertical ridges.

The distribution of this genus is restricted to the basin of the Amazon and the
rivers of Guyana. A key to the species has been furnished by Simpson (1914, p.
1212), but some of the species may be onl.y forms of others.

38. Hyria corrugata Lamarck (1819).

Hyria corrugata Lamarck and H. transversa Hupe, Simpson, 1900, pp. 868, 869;

1914, p. 1212, 1215.

Type-locality. — Unknown.

Other Localities. — Rio Solimoes (= middle Amazon) {Triplodon rugosum Spix,
Wagner, 1827, p. 35) ; Rio Yavari, Brazil (southern tributary of Amazon, forming a
boundary between Brazil and Peru) (Haas, 1916, p. 9, 47).

New Locality. — Sand-bar of Rio Tapajos, Santarem, Para, Brazil (J. D. Hase-
man coll., December 6-12, 1909). About fifty specimens, among them two fe-
males with soft parts.

562

MEMOIRS OF THE CARNEGIE MUSEUM.

Several other specimens are preserved in the Carnegie Museum, from the
Hartman, Smith, and Juny collections, but all are labelled ‘^Amazon River.”

Distribution. — According to Simpson Eastern Peru to Guiana; south through-
out Brazil. However, the few special localities known are all on the Amazon and
its tributaries.

It should be remarked in the first jilace that all mj^ sjiecimens undoubtedly
represent but one species, showing a great deal of variation, the various forms all
connected by intergrades.

Simpson descrilies this species as having a “subrhomboidal” shape and with
the beak-scul])ture covering only ])art of the surface. He retains H. transversa
Hupe (from Brazil) as a distinct species, which has “rhomboidal” shape, and only
a ‘‘few strong umbonal ridges.” Specimens from the Rio Tapajos at hand repre-
sent both forms, and pass gradually into each other, so that I am forced to regard
H. transversa as only an individual variation of H. corrugata.

The extreme develoimient of the lieak-sculjiture, covering the whole shell,
is found in //. rugosissinia Sowerliy. Some of my sjiecimens ap])roach this con-
dition, but none fully agree with Sowerby’s figure (XVH, Hyria, 1869, PI. 3, fig. 5),
so that this form is not represented in my material, and I cannot say whether it
is a good species or not.

Sowerby (/. c., PI. 2, fig. 3) distinguishes H. exasperata (British Guyana) from
H. corrugata (PI. 1, fig. 1), the former being longer and more compressed, the latter
shorter and more swollen. Sim])son unites these two; but among my material I
have no siiecimens which corresponds exactly to the corrugata of Sowerby. If the
two should be different, my siiecimens would fall under exasperata Sowerby (1869)
which undoubtedly is synonymous with Triplodon rugosum Sjiix (1829).

Frierson (1915, j). 363) has recently described H. amazonia. In outline this
easily falls within the range of my specimens of corrugata^ chiefly specimens with
less developed beak-scul])ture {H. transversa Hupe). But H. amazonia is decidedly
more swollen. Being founded upon a single specimen, it is impossible to decide
whether amazonia is a good species.

In shaiie my specimens vary greatly. They are all rather compressed, but
there is some variation in this respect. The greatest irregularity, however, is seen
in the outline and beak-sculpture. The wings of the shell are of various sizes,
chiefly the posterior one, which may be rather short, or drawn out, broader or
narrower, and may be more or less elevated. Sometimes the wings assume freakish
shapes, curving up or down, or being deflected laterally. Thus the whole shel}
is more or less elongated, and more broadly or more narrowly triangular. The

ortmann: south American naiades.

563

beak-sculpture is generally well developed, and covers all of the shell in young
specimens; but in older individuals it disappears near the lower margin, and some-
times it is poorly develojied, and present only for a short distance from the lieaks
(form transversa Hupe), but, as stated above, the latter siiecimens are connected
with the typical form by intergrades.

Anato7ny. — Two females are at hand, one of them gravid, and with eggs (col-
lected December 10)

The soft parts have been previously described and figured by myself (1911a,
pp. 108, 114, PI. 6, fig. 6, PI. 7, fig. 6).

Anal opening closed above; closed part about twice as long as the opening,
the latter slit-like, separated from the branchial o])ening by a mantle-connection.
Branchial opening with small papilhe, not closed in front. Palpi rather small,
siibtriangular, posterior margins not connected.

Gills of the usual shajie, the inner much wider anteriorly than the outer, its
anterior end close behind the palpi). Inner lamina of inner gill entirely connected
with abdominal sac. Non-marsupial gills with scattered interlaminar connections.
Marsupium of the female in the inner gill, restricted to the middle ])ortion, about
one-fourth of the gill at anterior end, and a little less than that at the posterior
end being non-marsupial. Interlaminar connections of marsupium crowded and
numerous, forming interruiited septa, and assuming in the most central part an
irregularly quincuncial (reticulate) arrangement.

Genus Prisodon Schumacher (1817).

Simpson, 1900, p. 869; 1914, p. 1216.

Closely allied to Hyria in the doubly alate shell, but differing in the absence
of beak-sculpture, and the strong develojiment of a sharp posterior ridge.

A key to the species has been given by Simpson (1914, ]). 1217).

This geniis is also characteristic of Guyana and the Amazon-drainage in Brazil.

39. Prisodon alatus (Sowerby) (1869).

Shells: Plate XL, figs. 1, 2, 3.

Hyria alata Sowerby, XVII, 1869, PI. 5, fig. 13.

Prisodon alatus Simpson, 1900, p. 871; 1914, p. 1220.

Type-locality. — Guyana.

The sliell of this gravid female ai^proaches the form transversa, with the beak-sculpture less
developed, but with the shape not very elongated. The other female is a normal corrugata, half-grown.

The connection of this gill with the palpi described previously is only accidental, and seen only
on one side of the larger female; on the other side, and also in the smaller female, the structure is normal.

5G4

MEMOIRS OF THE CARNEGIE MUSEUM.

New Locality. — Sand-bar of Rio Tapajos, Santarem, Para, Brazil (J. D, Hase-
man coll., December 6-12, 1909). About sixty specimens, all dead shells.

Distributioyi. — The discovery of this species, hitherto apparently only known
from Guyana, in the Amazon-drainage is interesting.

dlie shell is easily recognized by the relatively short triangular shape, and the
excessive develoinnent of the anterior and posterior wings. The outline varies a
good deal, lieing longer or shorter, and the lower posterior end of the shell may
be blunt or somewhat ])ointed. The oliesity is also variable. The wings are
more or less developed, and the ])osterior one may be longer or shorter, sometimes
as long as the rest of the shell, narrower or wider, pointed or blunt. It is directed
straight backwards (forming a straight continuation of the hinge-line), chiefly so
in young specimens; but in older ones it may be directed more or less upward, so
that the hinge-line becomes concave. Upon the posterior slope there are sometimes
a few (three to five) parallel folds or wrinkles, but most specimens lack these.

The pseudocardinals (PI. XL, figs. 2c, d) of this species are very long and com-
])ressed, the posterior of the right valve is longer than the anterior, and extends
far forwards, so as to be placed in its anterior part below the anterior. In the
left valve there are also two jiseudocardinals, the posterior shorter than the anterior,
and extending only with its anterior end below the anterior. The upper and inner
face of the posterior right pseudocardinal in its posterior part, and the space be-
tween the two teeth in the left valve, has distinct, parallel, and sub vertical ridges.
The lateral teeth (one in right, two in left valve) are only obliquely and granulately
corrugated, but do not possess parallel vertical ridges. Thus in the structure of
tlie })seudocardinals this species differs from other species of the genus (I have been
able to comjiare P. castelnaudi, obliquus, hroicnianus) , where these teeth are shorter,
and the iiosterior one in each valve is split into several parts, without developing
vertical ridges.

My specimens are of all sizes, but none of them reaches that of Sowerbj^’s
figure, which measures, from tip to tip of the wings, 115 mm., and from tip of an-
terior wing to lower ])osterior end of shell 115 mm.

Measurements.

T .1 /rp- c C Lpiieth (Anterior to

Length ( Tips of W.ngs.), ^

Heiglit.

^ Diameter.

Figured.

78 mm. 65 mm.

40 mm.

25.5 mm.

94 “ 1 69 “

43 “

29

93 “ ! 78 “

51 “

i 33

PI. XL, fig. I.

For obvious reasons (odd shape of shell) measurements could not be made here
in the usual way. It also should be noted that the wings of the largest specimens

ortmann: south American naiades.

5G5

are slightly broken off, and also in the two others they are not quite perfect. The
length from tip to tip of the wings is most nearly parallel to the hinge-line. The
height is measured vertically to the hinge-line, from the beaks to the lower margin.

40. Prisodon castelnaudi (Hupe) (1857).

Hyria castelnaudi Hupe, 1857, p. 81, PI. 16, fig. 1; Sowerby, XVII, 1869, PI. 4,

fig. 8.

Prisodon castelnaudi Simpson, 1900, p. 871; 1914, p. 1220.

Prisodon obliquus castelnaudi Von Ihering, 1910, p. 135, 137.

Type-locality. — Brazil.

Other Localities. — Rio Araguaya, Goj^az, Brazil (Von Ihering); Rio Chingu,
Para, Brazil (Von Ihering).

New Locality. — Rio Guapore, near Rio Sao Simao, Matto Grosso, Brazil
(J. D. Haseman coll., July 20, 1909). One young male with soft parts.

In addition, there is in the Carnegie Museum one larger specimen from the
Hartman collection, labelled “River Amazon.”

^distribution. — Positively known from the Amazon, two of the tributaries of
its lower part in Goyaz and Para, and from a tributary of the Madeira in Matto
Grosso.

M}^ specimens agree well with the account given by Simjison, but I should say
that the hinge-teeth are not vertically ridged. The posterior pseudocardinal in
either valve is cut up by deep fissures into accessory teeth, which are somewhat
radiating. The lateral teeth are slightly rugose.

Von Ihering regards this as a variety of P. obliquus Schumacher ( = avicularis
Lamarck), and this is quite possible.

Anatomy. — A young specimen with soft parts is at hand, probably a male,
since no marsupial structure can be seen.

Soft parts like those of Hyria. The palpi are slightly broader, and their
lower margins more curved. The anterior end of the inner gill is immediately
behind the palpi. Shape and structure of the gills normal.

Subfamily Mutelin^ (See above, p. 457).

General Remarks.

Shells of various shapes, subelliptical, subovate, subtrapezoidal, or more or
less rounded. Beak-sculpture absent, only in one case have subconcentric bars
been observed, which, however, may not be homologous to the real beak-sculpture
of other Naiades.

5G6

MEMOIRS OF THE CARNEGIE MUSEUM.

Hinge rarely with teeth, which, when present, are much reduced and consist
only of pseudocardinals or irregular teeth. Real laterals are never present.

Muscle-scars on inside of shell rather variable. Generally the anterior re-
tractor-scar is united with the adductor-scar, forming an upper continuation of
it (thus differing from the normal condition seen in the Hyriince). But in some
cases the retractor-scar is })artially or entirely isolated. Anterior jirotractor-scar
(piite varialile, isolated from adductor-scar or contiguous with it, or even confluent
with it. The iiosterior scars are rather uniform, agreeing with the Hyriince, the
retractor-scar forming an ipiiier process of the adductor-scar. Only in one case
(A nodontites erisiformis), are these two scars widely se])arated. Dorsal muscle-scars
mostly absent; if (rarel^b iiresent, only one inaj^ be found, or (in Leila) a row of
them. (In the Hyriince several are always jiresent.)

The pallial line is mostly siinjile and jiarallel to the margin; but in one genus
{Leila) it forms a shallow sinus posteriorly. This undoubtedly is connected with
the closing of the branchial opening in front, but unfortunately no soft parts of
this genus are at hand, so that jiarticulars cannot be given. Very often the pris-
matic liorder on the inside of the margin of the shell is unusually wide. -v

The ligamental sinus is comparatively large, larger than in the Hyriince; it
ma}' be broad and dee]i, but not shariily triangular {Iheringella), or deeji and sharply
triangular (in tlie other genera). In M ycetopocla it is shallow, but with a sharp
lower angle.

As to the characters of the soft parts see above (p. 457). But it is well to
])oint out here that the chief features in which this subfamily differs from the
Hyriince are found in the structure of the gills (text-fig. 3, i, o, p. 458) which have
well developed, solid sei)ta, moderately closely set, running parallel to the gill-
filaments (PI. XLVII, figs. 3, 4). In the female (PI. XLVII, figs. 35, 45; text-fig. 3f,
458), the sejita of the marsuiiial inner gill are stronger, but not distinctly more
crowded than in the non-marsupial gills, and have close to the outer lamina (primar}'
limb) a ridge on each side, projecting into the lumen of the water canal, incom-
pletely dividing the latter into two compartments (PI. XLVIII, figs. G, 75, 8).
The inner comi)artment, towards the inner lamina, assumes, when charged, the
function of an ovisac, containing the eggs or embryos (PI. XLVIII, fig. 75), which
do not stick closely together, and this compartment expands to a certain degree,
the corresponding section of the septa stretching out, while the outer compartment
(close to the other lamina) retains its shajie, and does not contain eggs, thus ap-
])arently serving as a secondary water-tube.

The size of the eggs is small, 0.07 to 0.09 mm. According to Von Ihering the

ortmann: south American naiades.

567

larval form is a lasidium (Size: 0.10 mm). It is a very singular circumstance
that I have not been able to find lasidia (or any other form of mature larvae) in
my material, although a good many gravid females of various species and genera
are at hand.

The Genera op the South American Mutelin^.

MutelincE are found in South America and in Africa. Unfortunately, among the
African forms, the anatomy of Spatha (including the subgenus Aspatharia) is
alone known (Ortmann, 1910, p. 39; 1918, p. 75); but this genus differs from most
South American Mutdince (Fossida, AJ onocondijlaxi, Anodontites) by the fact that
the anal opening is closed above, without forming a supra-anal opening. In the
South American genera this opening is open and not at all closed, with one exception,
Mycetopoda, where about the upper half of the anal is closed. In addition, in
Spatha, the inner gill is free from the abdominal sac, while in the South American
genera it is connected with it. All other characters are similar. Thus, although
closely allied to Spatha, the South American genera form a group by themselves,
and the similarity of Mycetopoda to Spatha in the anal opening apparently indicates ,
only parallelism of development, not genetic relationship.

It is hard to say which .group is more primitive, since of the two differing
characters, the one (anal opening) is more primitive in the American forms, the
other (inner lamina of inner gill) more primitive in the African Spatha. The
latter again is rather advanced in the shell, having no hinge-teeth, a condition
which is also found in most South American Muteliiue, lint not in all, for Iheringella,
Fossida, and Monocondykca have at least pseudocardinals.

There is no question that among the South American forms, these genera with
hinge-teeth should be regarded as more jirimitive. But a confirmation of this
cannot be found in the anatomy, the latter being alike in all of them. Possibly
the anatomy of Iheringella might furnish some enlightenment, but of this genus
the soft parts have never been observed.

The structure of the hinge might be expected to furnish evidence of the con-
nection of the Mutdince with the Hyriince. The very fact that there are genera
among the Mutdinm with hinge-teeth, indicates that the typical South American
forms (of the Anodontites-typa) are derived from forms with hinge-teeth. But
the structure of these hinge-teeth is rather peculiar. They are always in a rudi-
mentary condition, the laterals being missing, and the other teeth corresponding
to the pseudocardinals cannot be positively homologized with the pseudocardinals
of the Hyriince. Yet we are to assume on account of the many anatomical points

568

MEMOIRS OF THE CARNEGIE MUSEUM.

in common with the Hyriince that the Mutelinm are related to, and probably des-
cended from, the latter; but connecting links which undoubtedly stand between
these two subfamilies are as yet unknown, and only the presence among the Muteli-
nce of forms with hinge-teeth suggests that there once was a closer connection with
the Hyriince. According to our present knowledge, the two subfamilies are un-
doubtedly allied; but they are very sharply separated by anatomical as well as
shell-characters, and it is impossible to form an appropriate idea of their genetic
connection.

Simpson (1914, p. 1384 et seq.), in his family Mutelidce,, admits six South
American genera: Munocondykva, Ihermgella, Fossula, Leila, Anodontites, and
Mycetopoda. These are easily distinguished by shell-characters, which are tabu-
lated in the following key.

Key to the South American genera of Mutelin.e.

«!• Hinge with more or less developed pseiidocardinal teeth. Lunula short or almost absent, not much
produced in front of the beaks.

bi. Hinge-plate rather broad, with oblique, comjiressed or tubercular, alternating pseudocardinals,

two in each valve, the most anterior in the left valve Iheringella.

1)2. Hinge-plate narrow, with the pseudocardinals stumpy, squarish, or depressed, left valve never
with two distinct pseudocardinals.

Cl. Left valve with one, right with two stumpy, or squarish, pseudocardinals, which alternate,

the most anterior one being in the right valve Fossula.

C2. Either valve with only one pseudocardinal, which is generally depressed (spoon-shaped),

that of the left valve being the most anterior Monocondylcea.

0,2. Hinge without any pscudocardinal-teeth. Lunula elongated and much produced in front of the
beaks, forming a kind of anterior ligament.
bi. Pallial line without a sinus behind.

Cl. Shell rounded, subovate, or subtrapezoidal. Valves closed or somewhat gaping (wdien the
shell is elongated, the valves are closed). Anal opening entirely open. Branchial opening

not closed in front. Foot normal Anodontites.

C2. Shell elongated, gaping in front. Anal opening closed above. Branchial opening said to
be closed in front (this is doubtful). Foot very long, developed at its distal end into a

sort of button Mycetopoda.

b2. Pallial line with a sinus behind. Shell large, subovate, winged. Valves gaping. Branchial
opening said to be closed in front Leila.

With regard to the genetic connections of these genera, it is clear that we are
to regard the first three as the more primitive forms, and, judging from the shell,
Iheringella appears to be the most primitive, having the most complete hinge.
There is no doubt that of the others Anodontites is the simplest, but its connection
with the first three is obscure, since the shell-characters do not connect it more
closely with any one of them. Both Mycetopoda and Leila seem to have descended

ortmann: south American naiades.

569

from Anodontites, the former being characterized by a specialisation of the foot;
the, latter by a specialisation of the branchial opening. Mycetopoda is also peculiar

We can express this in the following

Leila

Anodontites

i

1

?

Iheringella, Fossula, Monocondylcea

The anatomy furnishes no additional help for the understanding of the phyl-
ogeny of these genera, except the points mentioned in the key. In fact, the anatomy
is so disappointingly similar in all of them, that it is practically useless. Only
in Mycetopoda and Leila have we been able to recognize higher stages of develop-
ment in the anatomy, but it should be mentioned that I have not been able to
verify this fact in Leila, of which I have not had the opportunity to examine soft
parts. The same is true of Iheringella. Of Fossula, Monocondylcea, and A710-
dontites, I know the anatomy very well, but no differentiations whatever have
been observed.

The Species of South American Mutelin,;E.

Genus Iheringella Pilsbry (1893).

Pilsbry, 1893, p. 30; Simpson, 1914, p. 1392.

Characterized by rather broad hinge-plate and compressed or stumpy pseudo-
cardinals, two in the right and two in the left valve, the anterior pseudocardinal
of the left valve the most anterior of all hinge-teeth. These teeth, however, are
rather variable.

This genus approaches the Hyrimce most closely in its hinge, but in the details
of its structure it is very different from any of them. It belongs to the La Plata-
drainage, and possibly also to that of the upper Amazon.

41. Iheringella balzani (Von Ihering) (1893).

Plagiodo7i halzani Von Ihering, 1893, p. 69, PI. 3, fig. 2.

Iheringella halzani Simpson, 1900, p. 914; 1914, p. 1359.

Type-locality. — Rio Paraguay, near mouth of the Rio Apa, Matto Grosso,
Brazil.

New Locality. — Rio Paraguay, San Luis de Caceres, Matto Grosso, Brazil
(J. D. Haseman coll.. May 25, 1909). Two complete shells and one left valve.

in having the anal opening closed above,
diagram :

Mycetopoda

570

MEMOIRS OF THE CARNEGIE MUSEUM.

Distribution. — Known only from the upper Rio Paraguay. Simpson also
gives “San Paulo, Brazil,” but 1 do not know upon what authority.

My specimens agree well with Von Ihering’s description and figure, but they
are larger. Von Ihering says that the color of the epidermis is dark olive or black-
ish. In our largest and smallest (half-shell) specimens the surface is much corroded
and worn, and brownish green. In the specimen of medium size, which is better
preserved, it is yellowish green, with a few scattered dark green spots upon the disk,
becoming more frequent near the beaks. Upon the posterior ridge, there are a
few fine, interrupted, dark green rays, and a broader ray runs over the posterior
slope.

In the two smaller specimens, the structure of the hinge corresponds entirely
with Von Ihering’s description. In the largest, however, the anterior tooth of
the left valve is obsolete, and so is the groove between the two teeth. The pos-
terior tooth is broad and slightly bifid. In consequence of this the groove of the
right valve is also broader and has a radial ridge in its bottom, and the anterior
tooth of the right valve is very small. I consider this an abnormality. In this
specimen there apparently is only one (bifid) tooth in the left valve, and two small
ones in the right, with an accessory ridge in the groove between them.

In all three of my specimens, the ligamental sinus is wider and shallower than
figured by Von Ihering, which may be due to their greater age.

Measurements.

Lengtli.

Height.

I>iameter.

Beaks.

34 Him.

20 nun. = 70 pr. ct. of L.

10 nun. =47 pr. ct. of L.

at 0 nun. = 20 pr. ct. of L.

40 “

31 “ =78

21 “ =53

11 “ =28

40 “

35 “ =70

23 “ =50

12 “ =20

32 “

27 “ =84

17 “ =53

= 30 “ (Von Ihering,

beaks from figure)

The measurements given by Simpson (1914, p. 1395) are entirely wrong, and
are those of Fossula balzani Von Ihering.

Genus Fossula Lea (1870).

Lea, Syn. 1870, p. 72, foot-note 1; Von Ihering, 1893, p. 62; Simpson, 1900, p. 914;
1914, p. 1396.

The distinguishing character of this genus is found in the hinge-teeth, of which
there are two in the right valve, enclosing between them one in the left valve. The
hinge-plate is narrow, the teeth are stumpy (not vertically compressed or spoon-like
as in M onocondylcBa) . There are traces of additional teeth and irregularities of
the hinge, which, however, are variable. The “cement processes” described by
Von Ihering are not always present.

ortmann; south American naiades.

571

A key to the three known species has lieen given by Von Ihering (1910, p. 115).
Two of the species are from the Parana-drainage, and one is from a coastal river
in eastern Brazil, the Rio Paraguassii in Bahia.

42. Fossula fossiculifera (D’Orbigny) (1835).

Section of gills: Plate XL VIII, fig. 6.

Monocondylcea fossiculifera D’Orbigny, 1843, p. 614, PI. 80, figs. 5-7.

Unio fossiculif eras Sowerby, XVI, 1868, PI. 96, fig. 521.

Fossula fossiculifera Von Ihering, 1893, p. 64, PI. 3, fig. 2; Nehring, 1893, p. 164;
Simpson, 1900, p. 914; Von Ihering, 1910, p. 115; Simpson, 1914, p. 1396.
Type-locality . — Rio Parana, Iribiiciia, above Corrientes, Argentina.

Other Localities. — Rio Parana and lower Rio Tiete, Sao Paulo, Brazil (Von
Ihering); Rio Piracicaba, Piracicaba, Sao Paulo, Brazil (Von Ihering) (Nehring).

Localities Represented in the Carnegie Museum. — Rio Piracicaba, Piracicaba,
Sao Paulo, Brazil (Von Ihering donor). One specimen. Rio Tiete, Salto de
Avanhandava, Sao Paulo, Brazil (J. D. Haseman coll., September 15, 1908). One
specimen. Rio Tiete, Salto das Cruzes, Sao Paulo, Brazil (J. D. Haseman coll.,
September 22, 1908). One female with soft parts. Rio Tiete, Itapura, Sao
Paulo, Brazil (J. D. Haseman coll., September 28, 1908). One female with soft
parts.

Distribution. — Rio Parana and its tributaries from Argentina to Sao Paulo,
Brazil.

This species has been well figured liy D’Orbigny and Von Ihering, and has
been well described by the latter. No further details need to be supplied.

Measukements.

Sex,

Length.

Height.

Diameter.

Beaks.

Salto das Cruzes

9

56 mm.

44 mm

= 78 pr. ct. of L.

27 mm

= 48 pr. ct. of L.

at 21 mm

= 38 pr. ct. of L.

Itapura

9

74 “

63 “

= 85

38 “

= 51

23 ‘‘

=31

Avanhandava

79 “

62 “

=78

38 “

= 48

25 “

= 32

Piracicaba

79 “

64 “

= 81

39 “

= 49

26 “

= .33

(Von Ihering)

74 “

63 “

= 85

38 “

=51

24 “

=32

According to Von Ihering the height varies from 80 to 85 pr. ct., while in D’Or-'
bigny’s specimen it is 77 pr. ct.

Anatoyny. — Partially described by Von Ihering (1893, p. 65) for the male. I
am able to supplement this from two barren females at hand.

Anal opening entirely open, very large, with smooth inner edge, very faintly
crenulated in the lower part, separated by a solid mantle-connection from the

572

MEMOIRS OF THE CARNEGIE MUSEUM.

branchial opening, which has small papillae on the inner edge. These papillae
extend rather far forward, decreasing in size, and disappearing gradually. Palpi
moderately large and broad, semicircular, posteriorly with a short truncation
forming the posterior margins, which are not connected.

Gills rather wide, the inner the wider, chiefly anteriorly, its anterior end
immediately behind the palpi; that of the outer gill at the highest point of the
attachment-line of the mantle, so that the lower margin of the outer gill is curved,
the gill narrowing considerably in front, while the inner is not much narrower in
front, and has the lower margin rather straight. Inner lamina of inner gill entirely
connected with abdominal sac. Both gills with well developed, strong septa. The
non-marsupial outer gill has the septa alternately stronger and thinner, but the
alternation is irregular. The marsupimn is in the inner gill of the female. Here
the septa are more uniform and very strong, but hardly more closely set than in
the non-marsupial gill. Where the septa are inserted at the outer limb, they have
a swelling on each side, which forms a vertical ridge projecting into the lumen of
the water-tubes (See PI. XL VIII, fig. 6). This ridge is less developed in my younger
female, yet perfectly distinct. The sei)ta near the anterior and posterior end of
the inner gill do not have marsupial structure, but the marsupial part is very large,
occupying nearly the whole gill.

Genus Monocondyla^a D’Orbigny (1835).

D’Orbigny, 1835, p. 37; 1843, p. Gil; Simpson, 1900, p. 910; 1914, p. 1384.

Normally each valve has only one iiseudocardinal tooth, that of the left valve
being more anterior. The teeth are more or less depressed (spoon-shaped). Traces
of additional teeth may be present, but they are insignificant and variable.

A key for the species has been given by Simpson (1914, p. 1385), which, how-
ever, is artificial and unsatisfactory, and the essential characters apparently have
been misunderstood. Aloreover, I believe that M. guarayana D’Orbigny (1843,
PI. 68, figs. 4-7) does not belong here, but jirobably to Ihei'ingella.

The specimens before me all have a cloth-like epidermis, with crowded, ir-
regular and anastomosing, concentric lines, which are lamellarly elevated. But
very often these fine lamella} are abraded, so that the surface appears smooth.
Nevertheless the lamellae are generally i)reserved in some part of the shell, chiefly
on the posterior slope and near the margins, and, even when abraded, they can be
easily noticed as fine lines. The species at hand, which have been treated by Simp-
son have been called ‘‘smooth or lightly concentricalD striate.” He distinguishes
them according to the outline, orbicular, obovate, elliptical, subrhomboidal, or
subquadrate, but it is extremely hard to draw a line between these terms.

ortmann: south American naiades.

573

The following key is intended only for the distinction of the forms at hand.

Key to the Species of Monocondyl^a.

ai. Shell small, or of medium size (at the utmost 50 to GO mm. long). Lamellar striie of epidermis well
developed, but sometimes abraded. Color of epidermis greenish black or brown.
hi. Shell more or less oblique, subtrapezoidal, or angularly suliorbicular, with a well developed
posterior upper angle, high behind, narrower in front. Color of epidermis greenish black.
Prismatic zone on inside of margin of unequal width, very wide along anterior portion of
lower margin.

Cl. Shell angularly suborbicular, rather high and short, height about 80 pr. ct. of length. Sides
of disk not llattenetl.

di. Shell compressed, diameter 50 iir. ct. or less of length. Beaks not much elevated.

M. lentiformis.

df Shell more swollen, diameter 58 to 59 pr. ct. of length. Beaks more inflated and

elevated M. paraguayana.

C2. Shell subtrapezoidal, more elongated, height about 70 pr. ct. of length. Sides of disk more
or less flattened.

di. Shell more swollen, beaks more prominent, hinge-line rather strongly incurved under

the beaks M. minuana.

d-i. Shell less swollen, beaks less prominent, hinge-line less incurved under the beaks.

M. minuana parchappi.

62. Shell very little, or not at all oblique, subovate or subelliptical in outline, with the posterior
upper angle rounded and poorly developed. Obesity of shell very great, and beaks much
swollen and inflated. Color of epidermis brown. Prismatic zone on inside of margin narrow

and of nearly uniform width M. obesa.

Ui. Shell large (about 100 mm. long). laimellar striae of epidermis poorly developed upon the disk,
distinct only near the margins. Color of ejiidermis from yellowish olive upon the disk and toward
the beaks to dark brown toward the margins M. hollandi.

The species of this genus cover a rather large range in South America being
found all over the La Plata-drainage, in the Amazon-drainage in Bolivia and
Brazil, in the Rio San Francisco in eastern Brazil, and also in coastal streams in
southern Brazil (Rio Grande do Sul).

43. Monocondyl^a lentiformis Lea (1866).

Anatormy of gills: Plate XLVII, fig. 3.

M onocojidylcea lentiformis Lea, Obs., XII, 1869, PL 36, fig. 86; Pilsbry & Rush,
1896, p. 81; Simpson, 1900, p. 912; 1914, p. 1392; Corsi, 1901, p. 452; Haasi
1916, pp. 25, 54.

Aplodon lentiformis Von Ihering, 1893, p. 67; Nehring, 1893, p. 164.
Type-locality. — South America.

Other Localities. — Rio Piracicaba, Piracicaba, Sao Paulo, Brazil (Von Ihering)
(Nehring); Rio Uruguay, Salto Oriental, Uruguay (Haas); Rio de la Plata, Colonia,
Uruguay (Pilsbry & Rush).

574

MEMOIRS OF THE CARNEGIE MUSEUM.

New Localities. — Rio Uruguay (in mud), Uruguayana, Rio Grande do Sul,
Brazil (J. D. Hasenian coll., February 5, 1909). Four specimens with soft parts,
males and females. Rio Cacequy, Gacequy, Rio Grande do Sul, Brazil (tributary
to Rio Ibicuhy, to Uruguay) (J. D. Haseman coll., February 2, 1909). Two speci-
mens with soft iiarts, male and female. Rio Jacuhy, Gachoeira, Rio Grande do
Sul, Brazil (tributary to Rio Guahyba) (J. D. Haseman coll., January 26, 1909).
Two specimens.

Distribution. — Known from the Rio de la Plata, and up the Uruguay, and from
the upper Parana drainage in Sao Paulo. The species has not been found in the
lower and middle Parana, although it should be expected there. In addition it is
found, as our specimens show, in the drainage of a coastal stream in Rio Grande do
Sid (Guahyba). This is a remarkable fact, but it should be borne in nund that the
head-waters of the Ibicuhy (Uruguay drainage) and those of the Jacuhy (Guahyba-
drainage) closely interlock, and that stream piracy in this region is quite probable.

Characters of the Shell. — Shell rather small (maximum length 44 mm.), moder-
ately thick, outline more or less subcircular or subpolygonal. Height 77 to 88 pr.
ct. of length. Valves very little gaping in front, sometimes almost closed. Dorsal
margin straight behind the beaks, or gently convex, much lower in front of the
beaks, and straight here or even somewhat concave, passing in a curve or a very
blunt angle into the anterior margin. Posteriorly the dorsal margin passes in
a distinct, or indistinct, blunt angle into the posterior margin. The latter descends
obliquely, and is straight, rarely slightly convex or concave. It passes in a curve
into the {losterior part of the lower margin, forming the blunt posterior end of the
shell. The lower margin is curved, the strongest part of the curve and the lowest
lioint of the shell is situated at 54 to 60 pr. ct. of the length from anterior end, and
is thus immediately liehind the middle. From this point the lower margin curves
up rather strongly in the posterior part. In the anterior portion it is very gently
curved, or almost straight, and finally curves up sharply into the anterior margin.
Thus the anterior end of the shell appears narrower than the posterior, and the
shell has the shape of an irregular pentagon.

VaKcs very gently convex, rather compressed, greatest convexity behind the
middle of the shell, without a distinct posterior ridge. Sides not flattened. Toward
the anterior and })osterior end the shell is somewhat comiiressed. Diameter 41
to 51 pr. ct. of the length. Beaks little swollen, and very little prominent, located
at 31 to 39 pr. ct. of the length. Lunula narrow and short.

Ei)idermis dull, not shining, with very crowded, irregular concentric lines,
which form fine lamellae, and give a cloth-like appearance to the surface. But

ortmann: south American naiades.

575

these lamellie may be worn off in i)art, and the surface may be smoother. No
distinct radial sculpture is visible, but there may be a trace of a radial rib upon the
posterior slope. Color of epidermis blackish green, without color-markings, ex-
cept traces of one or two dark rays upon the posterior slojie.

Hinge-line straight behind the beaks. In front it is curved down more or
less suddenly, and carries a single pseiidocardinal tooth in each valve. The tooth
of the left valve stands in front of, and somewhat below, that of the right valve.
Both teeth are vertically depressed, somewhat spoon-shaped, and project under
the edge of the ojiposing valves into corresiionding shallow grooves. Ligamental
sinus triangular, not dee]ier than wide; its anterioi- margin more or less oblique to
the hinge-line.

Nacre very iridescent, whitish, but generally with salmon, pink, or pale purple
stains, chiefly in the cavity of the shell. Irregular radiating lines present, but
only near the margin. Nacreless (prismatic) zone of irregular width, narrow be-
hind, suddenly increasing in width at the lower point of the lower margin, and re-
maining wide to near the anterior margin. Greatest width of this zone over half
the distance of mantle-line from margin. Color of nacreless zone dull olive, gray-
ish, or grayish red.

Cavity of shell and beaks shallow. Anterior adductor-scar well marked, ovate.
Anterior retractor-scar small, but distinct, separated from or connected with the
adductor-scar, lying in the left valve on the base of the jiseudocardinal tooth. An-
terior protractor-scar connected with adductor-scar. Posterior adductor-scar
faint, ovate; posterior retractor-scar foi'ining a tilangular up])er pi-ojection of
adductor-scar. No dorsal scars. Pallial line subconcentric to the margin.

Measurements.

Localities.

No.

Sex.l Leu

1

Height.

Diameter.

Beaks.

Greatest Height.

Cachoeira . . .

1

34 nun. 27 nim. =79% of L.

14 nun. =41 % of L.

at 1 1

mm. =32% of L.

at 20.5 mm. =00% of L.

Uruguay ana .

1

38

“ 31.5

“ =83 “

11

X

15

“ =39 “

22 “ =58 “

Cacequy ....

2

d'

39

“ I30

“ =77 “

10 “ =41 “

13

“ =33 “

23 “ =59 “

Do

1

9

39

“ '31

“ =79 “

18 “ =40 “

13

“ =33 “

23 “ =59 “

Uruguayana .

3

9

39

“ ,34.5

“ =88 “

19.5 “ =50 “

14

“ =30 “

21 “ =.54 “

Do.

2

41

“ '33.5

“ =82 “

18 “ =44 “

13

“ =32 “

24 “ =59 “

Do.

4

9

42

“ 30

“ =80 “

20.5 “ =49 “

15

“ =30 “

23 “ =.55 “

Cachoeira . . .

3

?

43

“ |34

“ =79 “

19 “ =44 “

13.5

“ =31 “

25 “ =.58 “

Remarks. — These measurements agree well with those given by Von Ihering
and Nehring. According to Von Ihering the height ranges from 80 to 84 pr. ct. of
length, the diameter from 46 to 51 pr. ct., while Nehring gives for the height 77 to 82
pr. ct., the diameter 46 to 49 pr. ct. Lea’s original figure shows, according to
Von Ihering, 80 pr. ct. for the height, and 44 pr. ct. for the diameter. The speci-

576

MEMOIRS OP THE CARNEGIE MUSEUM.

mens from Cachoeira (Guahyba-drainage) entirely correspond with the others
in these respects, and I cannot distinguish them by any other characters.

The chief features of this species are the subcircular, or rather rounded-pen-
tagonal outline, the great height of the shell, the location of the greatest height
close behind the middle of the shell, and the flatness of the valves. If properly
l)laced, with the ligament horizontal, it is seen that the shell is distinctly oblique,
and that the anterior end is narrower.

Anatomy. — Three males and three barren females are at hand.

Von Ihering (1893, p. 69) has mentioned a number of anatomical characters,
but his account is not complete.

Anal opening entirely open, large, its inner edge with distinct crenulations
in the lower part, otherwise smooth, separated from the branchial opening by
a mantle-connection. Inner edge of branchial opening with distinct, but small,
papilla3. Palpi moderately large, semicircular, briefly truncated behind, the pos-
terior margins not connected.

Gills (PL XLVII, figs. 3a, h) of medium width, the inner the wider, chiefly in
front; the outer narrowing anteriorly, its anterior end near the highest point of
the mantle-attachment-line. The inner gill very little narrowed in front, beginning
immediately behind the palpi. Inner lamina of inner gill entirely connected with
abdominal sac.

Gills with well developed septa. Those of the male (PI. XLVII, fig. 3a) and
of the outer gill of the female alternately stronger and weaker. This alternation,
however, is not very distinct, and is chiefly seen in the middle of the gill. In
the female (PI. XLVII, fig. 35) the inner gill is marsupial, with the septa more
uniform and stronger, but hardly more crowded. At the point of union with the
outer lamina, the usual swelling is present, indicating the ridges projecting into
the water tulies. The most anterior and most posterior extremity of this gill
has not the marsupial structure, but the marsupium occupies nearly the whole
gill.

In the siiecimen sectioned, the swellings of the septa are located nearly in
the middle. However, the part of the septa from the swelling toward the inner
lamina is thicker, and has more strongly developed epithelium, indicating that
this is the part which stretches out, when gravid. The condition seen in this
s]iecimen undoubtedly is due in part to its barren character, in jiart to the state
of ]ireservation.

ortmann: south American naiades.

577

44. Monocondyl^a paraguayana D’Orbigny (1835).

Monocondyloea paraguayana D’Orbigny, 1843, p. 612, PL 70, figs. 5-7; Simpson,

1900, p. 911 (in part); 1914, p. 1387.

JJnio paraguayana Sowerby, XVI, 1866, PI. 52, fig. 273.

Type-locality . — Rio Parana, Italy, near Corrientes, Argentina.

Other Locality. — Rio Batel, Province Corrientes, Argentina.

New Locality. — Rio Uruguay (in mud), Uruguayana, Rio Grande do Sul,
Brazil (J. D. Haseman coll., February 5, 1909). One female with soft parts.

DiMribution. — Known from Rio Parana and its tributary, the Rio Batel in
Argentina, and from the Rio Uruguay.

Characters of Shell. — Shell moderately thick, and of medium size (maximum
length 59 mm.). Outline briefly subtrapezoidal, rather high (height 77 to 81 pr. ct.
of length), distinctly oblique, but approaching an angularly suliorbicular shape.
Valves in front gaping very little. Dorsal margin behind the beaks slightly convex,
much lower in front of the beaks, where it is distinctly concave, and passes into the
anterior margin in a blunt, but distinct, angle. Posteriorly the dorsal margin passes
into the posterior margin in a rather sharp curve, forming a blunt angle. Posterior
margin descending obliquely, but very steejily, almost straight, and curving into the
lower margin, forming with it the broadly rounded iiosterior end of the shell.
Lower margin very little curved in the anterior part and sloping upward ; posteriorly
more distinctly curved. Its lowest point (and the greatest height of the shell) at
about 57 pr. ct. of the shell (60 pr. ct. according to D’Orbigny ’s figure). In front,
the lower margin curves up into the anterior margin. The anterior end of the
shell is narrower than the posterior, thus producing the oblique shape.

Valves very convex, chiefly so in the anterior part and backward to the pos-
terior ridge, and without any flattening upon the sides. Behind the posterior
ridge the shell is strongly compressed, so that the posterior slope and the upper
posterior angle appear almost alate. The most extreme anterior end of the shell
is also slightly compressed. Diameter 59 pr. ct. of the length (58 pr. ct. according
to D’Orbigny). Beaks inflated and swollen, incurved, projecting over the lunula,
but only slightly elevated above the posterior part of the upper margin. Lunula
distinct, short, triangular, about half as wide as long.

Epidermis dull, with crowded, irregular, concentric lines, which frequently
become lamellar: probably, where not lamellar, they are worn off, and originally
the whole epidermis was cloth-like. No radial sculpture visible. A radial rib
upon the posterior slope, but rather indistinct (much less distinct than in D’Or-

578

MEMOIRS OF THE CARNEGIE MUSEUM.

bi^iiy’s figure, and hardly forming a point and an emargination on the posterior
margin, as in this figure). Color of epidermis dark grayish green to blackish, the
grayish green shade due to the iireservation of the lamellae. A rather distinct
black radial ray u])on the rib of the posterior slope, and faint traces of additional
rays in front of and behind it.

Hinge gently curved behind the beaks. Under the beaks it is sharply curved
down, and in front of them it curves up again, thus becoming concave. Each
valve has a single ]iseiidocardinal tooth of the typical shape; that of the left valve
anterior to, and a little lielow, that of the right; both triangular and depressed,
and projecting under the margin of the ojiposite valve, where they fit into shallow
grooves. Ligamental sinus triangular, wider than dee]), its anterior margin oblique
to the hinge-line.

Nacre very iridescent, whitish, with a faint salmon blush in the cavity of
the shell, and jiurple and greenish reflections toward the margins. Irregular
radiating lines are present toward the margin. Prismatic zone of unequal width,
rather narrow behind, widening quite suddenly at the lowest ])oint of the shell,
and remaining wide along the anterior ascending jiart of the lower margin, narrowing
again at the anterior end. Color of prismatic zone grayish green.

Cavity of shell and beaks moderate. Anterior adductor-scar dee]), elliptical.
Anterior retractor-scar small, at the ii])])er end of the adductor-scar, distinctly
isolated in left valve, nai-rowly connected with it in the right. Anterior ])rotractor-
scar small and united with adductor-scar. Posterior adductor-scar faint, subovate,
with an u])])er triangular projection formed by the posterior retractor-scar. No
dorsal scars. Pallial line subconcentric to the margin.

]\rE.A.SUKEMENTS.

Sex.

Length.

Height. Diameter.

Ifeaks.

Greatest Height.

9

.'53 mm.

43 mm. =81 pr. cl. of L. Ill mm. =.59 pr. ct. of L.

at 16 mm. =30 pr. c

t. of L.

at 30 mm. =57 pr. ct. of L.

Remarks. — In general outline this species is much like M. lentiformis, as is
shown by the ])roportion of height to length and the location of the greatest height
of the shell. It markedly differs, however, in the much more swollen shell (diameter
59 ])r. ct. as against 41 to 51 ])r. ct. in lentiformis) , and the more inflated beaks.
In consequence of the very convex disk, the ])osterior slo])e appears much more
compressed and alate.

Anatomy. — The s])ecimen at hand for examination is a barren female.

The structure is absolutely identical with that of M. lentiformis, and nothing
is to be added, except that the crenulations of the lower ])art of the anal opening

ortmann: south American naiades.

579

are less distinct, and that the palpi appear a little larger, with longer posterior
margins, which are connected at the base. These differences, however, may be
due to preservation.

The septa of the inner marsupial gill also have the characteristic swelling,
not so close to their insertion with the outer lamina, but more toward the middle
of the septum.

45. Monocondyl^a minuana D’Orbigny (1835).

Section of gills: Plate XLVITI, fig. 7.

Monocondyloea minuana D’Orbigny, 1843, p. 612, PI. 70, figs. 8-10; Corsi, 1901,
p. 452, fig. 34; Simpson, 1914, p. 1388.

Unio minuanus Sowerby, XVI, 1868, PI. 91, fig. 497.

Monocondylcea pazii Lea, Obs., XII, 1869, PI. 36, fig. 88; Pilsbry & Rush, 1896,

p. 81.

Monocondylcea paraguayana Simpson, 1900, p. 911 (?iro parte).

Type-locality . — Canelon Grande and del Rosario, Banda Oriental, Uruguay
(Arroyo Grande flows North to Rio Negro; Arroyo Rosario flows South to Rio de la
Plata).

Other Localities. — Arroyo de las Vacas, Uruguay (Corsi); Rio de la Plata,
Colonia, Uruguay (Pilsbry & Rush, pazii).

New Localities. — Rio Uruguay, (in mud) Urugiiayana, Rio Grande do Sul,
Brazil (J. D. Haseman coll., February 5, 1909). Five specimens with soft parts,
males and females. Rio Jacuhy, Cachoeira, Rio Grande do Sul, Brazil (J. D.
Haseman coll., January 26, 1909). Five specimens with soft parts, males and
females.

Distribution. — Rio de la Plata and its tributaries in the Banda Oriental; Rio
Uruguay and Rio Negro drainages; and in Guahyl^a drainage in Rio Grande do Sul.

Characters of the Shell. — Shell moderately thick, rather small (maximum length
55 mm.). Outline subtrapezoidal, strongly oblique, rather elongated (height 07
to 72 pr. ct. of length). Valves very slightly gaping at anterior end. Dorsal
margin straight behind the beaks, or gently convex, much lower in front of beaks,
where it may be straight or somewhat concave, forming with the anterior margin a
more or less distinct, obtuse angle. Posteriorly the dorsal margin passes into the
posterior margin in a more or less distinctly obtuse angle. Posterior margin ob-
liquely descending, straight, or gently curved, passing into the lower margin in
a strong curve, which forms the rounded lower posterior end of the shell, not
much elevated above the base-line. Lower margin in its middle and anterior part

580

MEMOIRS OF THE CARNEGIE MUSEUM.

very gently curved or almost straight, sloping upward; the lowest point located
rather far backward (at 66 to 77 pr. ct. of the length). Anteriorly and posteriorly
the lower margin curves up into the anterior and posterior margins. Anterior
end of shell markedly narrower than the posterior.

Valves very convex, but distinctly flattened upon the sides of the disk, which
flattening may even become a shallow dejiression. Greatest convexity in front
and behind the flat area, and the posterior convexity, forming the rounded pos-
terior ridge, is the stronger. Posterior slope somewhat compressed and elevated,
but hardly alate. At its anterior end the shell is also a little comiiressed. Diameter
50 to 56 ]ir. ct. of length, but sometimes falling below 50 pr. ct., and such specimens
approach the var. parchappi. Beaks more or less swollen, inflated, and incurved,
prominent over the lunula, but only little elevated above the jiosterior part of
the iijiper margin. Lunula distinct, short, elongated triangular, variable, narrower
in young, broader in old specimens, where it may be half as wide as long.

Epidermis dull, cloth-like. Fine and crowded, anastomosing, concentric
striae are elevated as fine lamellae, but often they are abraded. No radial sculpture
visible, and there is hardly a trace of a radial ridge upon the posterior slope. Color
of epidermis dark grayish green, without any trace of color-markings, except oc-
casionally a mere indication of a dark ray upon the posterior slope.

Hinge straight, or gently curved, in its posterior part. Under the beaks it
curves down more or less suddenly, and then becomes straight again, or even some-
what concave. The curvature of the hinge-line is quite variable, and corresponds
to the vaiying degree of the develoinnent of the beaks. Pseudocardinal teeth
as in M. parnyuayana. Ligamental sinus triangular, wider than deep, with its
anterior margin oblique to the hinge-line.

Nacre vei\y iridescent, whitish, with greenish and purplish tints, chiefly pos-
teriorly. Irregular radiating lines jiresent, but faint in old specimens. Pris-
matic zone of unequal width, rather narrow behind, suddently widening at the
lower jioint of the lower margin, and remaining wide to near the anterior margin.
However this widening is not so strongly pronounced as in M. paraguayana, and
is also individuality varialile. Color of iirismatic zone greenish gray.

Cavity of shell and lieaks moderate. Anterior adductor-scar sharply and
rather dee])ly impressed, subelliiitical; anterior retractor-scar small, connected with,
or seiiarated from, adductor-scar; anterior protractor-scar connected with adductor-
scar. Posterior adductor-scar distinct, but less strongly impressed; scar of the
]iosterior retractor forming a short, triangular upper process of it, which, in a few
siieciniens, is jiartly separated from it. No dorsal scars. Pallial line distinct,
subconcentric to the margin.

ortmann: south American naiades.

581

Measurements.

Localities.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Greatest Height.

Uruguay-
ana

1

d'

24

mm.

17 mm. =71% of L.

12 mm. =50% of L.

at 9 mm. =38% of L.

at 18.5

mm. =77% of L.

Do. . .

3

9

35

25

“ =71 “

17.5

“ =50 “

12 “ =34

25

“ =71 “

Do. . .

C

9

39

28

“ =72 “

22

“ =56 “

14 “ =36

27

“ =69 “

Do. . .

5

d

41

28

“ =68 “

21

“ =51 “

14 “ =34

27

“ =66 “

Do. . .

7

d

48

33

“ =69 “

24

“ =50

15 “ =31

33

“ =69 “

Cachoeira . .

1

d

28

19.5

“ =70 “

12.5

“ =45 “

9.5 “ =34

18.5

“ =66 “

Do. . .

2

9

34.5

23

“ =67 “

16.5

“ =48 “

■ 11 “ =32

23

“ =67 “

Do. . .

X

?

36

27

“ =75 “

16.5

“ =46 “

11 “ =31

25

“ =69 “

Do. . .

3

9

43

30

“ =70 “

22

“ =51 “

13 “ =30

29

“ =67 “

Do. ..

4

9

44

30.5

“ =69 “

21.5

“ =49 “

12 “ =27

29

“ =66 “

Remarks. — The measurements for minuana given by D’Orbigny in the text and
those taken from his figures do not agree, but according to the latter the height
is about 64 to 67 pr. ct. of length, and the diameter about 50 pr. ct. Our specimens
from Cachoeira have, on the average, a smaller diameter (falling as low as 45 pr.
ct.), and in this respect they are transitional toward the variety parchappi; but in
other characters (beaks and hinge-line) they agree better with typical minuana.

M. pazi Lea is apparently the same species. Simpson unites it with parchappi,
and it indeed approaches the latter in obesity, having a diameter of 46 pr. ct.,
{parchappi of D’Orbigny has 43 pr. ct.), but here again the beaks and the curvature
of the hinge-line are more like minuana, and for this reason I place it here.

Just because such intergrades do exist, I regard parchappi as a variety of
minuana (See below). Where the line between the two forms should be drawn re-
mains doubtful, and naturally it could not be expected that there is a sharp line. I
have named as miuana my specimens with the diameter of 45 pr. ct. and over,
because their beaks and hinge-line are more like those of minuana, but this procedure
possibly may require modification, when more material is studied.

Anatomy. — I have examined the soft parts of six males, and of five females.
Two of the latter, collected February 5 (Uruguayana), were gravid.

Lea (Obs. XII, 1869, p. 273) has described the soft parts of M. pazi, and, as
far as it goes, this description agrees with our specimens.

The structure of the soft parts is essentially like that of the foregoing species
{lentiformis and paraguayana) . It should be remarked that the palpi are rather
. large, with a somewhat longer posterior truncation, and the posterior margins are
connected at the base. Anal opening practically smooth. Papillie of the branchial
opening very small.

In the gravid female the whole inner gill is charged, with exception of the
outermost extremities. The swelling of the marsupium is very moderate, and in a
cross-section (PI. XLVIII, fig. 7b) it is seen that only the part of the septa, extend-

582

MEMOIRS OF THE CARNEGIE MUSEUM.

ing from the swellings, toward the inner lamina (secondary limb) stretch out, and
that the egg-masses are located only in this ])art of the water-tubes, while the
])art toward the outer lamina (primary limb) does not contain eggs, and forms
what should be called secondary water-tubes. In the sterile female the swelling
of the septa (vertical ridges projecting into the water tubes) are located as usual.
In the cross-section of the gills of the male (PI. XLVIII, fig. la) it is seen, chiefly
in the inner gill, that the alternation of stronger and weaker septa is due to the
alternating presence or absence of a larger blood-vessel at the point where the
septum connects with the ])rimary limb. In the marsupial gill of the female this
is obscured in the direct view by the development of the ridges, so that the septa
appear more uniform.

The contents of the charged marsupium consist of small globular embryos in
an early stage of development. No lasidia were seen. Von Ihering (1891, p.
480) mentions the eggs of Aplodon pazi. They are small, 0.075 mm. in diameter.

45a. Monocondyl^a minuana parchappi (D’Orbigny) (1835).
Mo7iocondylcea parchappi D’Orbigny, 1843, p. G15, PI. 68, figs. 1-3; Simpson,

1900, p. 911; 1914, p. 1386; Haas, 1916, pp. 24, 54.

Type-locality . — Rio Parana, Italy, Province Corrientes, Argentina.

Other Locality. — Rio Uruguay, Salto Oriental, Uruguay (Haas).

New Locedity. — Rio Jacuhy, Cachoeira, Rio Grande do Sul, Brazil (J. D. Hase-
man coll., January 26, 1909). One specimen, male with soft parts.

Distributiori. — Rio Parana and Rio Uruguay, and also Guahyba-drainage.
The distribution resemliles that of M. minuana.

D’Orbigny has already iiointed out the close resemblance of this form to M.
niinuaria, but gives as differentiating characters the greater compression of the
shell, the less elevated and less incurved beaks, the absence of a lunula, and the
rose-colored nacre. We may dismiss the last two characters as unim]iortant,
since the lunula is variable and generally less develojied in forms with lower beaks;
and since reddish nacre is found as an individual variation in other siiecies of
Monocondyloea. But the greater compression of the shell and the feebler develop-
ment of the lieaks is striking. In D’Orbigny ’s figure, as well as in my specimen,
the latter character is connected with a straighter hinge-line, which is much less
incurved below the beaks.

We have seen that specimens of minuana sometimes approach parchappi in
the less ])ronounced obesity. The same may be said of the inflation of the beaks
and the curvature of the hinge-line. However, none of my S]Decimens I’ecorded as

ortmann: south American naiades.

583

minuana from Caclioeira have the beaks as low and the hinge-line as straight as
the specimen placed under yarcliappi. But there is no question that the}^ incline
in this direction, and that beaks and hinge-line are quite variable in minuana.
Thus these two forms should be regarded as varieties of one species, actually con-
nected by intergrades.

Measurements.

Sex.

Length.

Height. 1 Diameter. ! Beak.s.

Greatest Height.

cf

41 mni.

29 mm. =71 pr. ct. of L. 17 mm. =41 pr. ct. of L. at 12 mm. =29 pr. ct. of L.

at 26 mm. =63 pr. ct. of L.

According to D’Orbigny, parchappi has a height of 64 pr. ct. and a diameter of
43 pr. ct., so that length and diameter are a little greater than in my specimen.

Anatomy. — The specimen at hand is a male according to the soft parts. The
structure is absolutely identical with that of the males of AI. ‘minuana.

46. Monocondyl^a obesa Ortmann, sp. nov.

Shells: Plate XL, figs. 4, 5, 6.

Type-locality. — Rio Tapajos, Santarem, Para, Brazil (J. D. Haseman coll.,
December 6-12, 1919). Type-set: Carn.'Mus. Cat. No. 61.5850. Seventeen com-
plete shells and a number of odd valves.

Characters of Shell. — Shell small to medium in size (maximum length 57 mm.),
moderately thick, outline briefly subelliptical, subovate, or subrotund (when
young), hardly oblique. Height 75 to 86 pr. ct. of the length. Valves not gaping.
Dorsal margin behind the beaks gently curved or almost straight, subconcave and
very short in front of the beaks, passing gradually or at an indistinct angle into
the anterior margin. Posteriorly the dorsal margin passes by a blunt, indistinct
angle, or almost gradually, into the posterior margin, which descends obliquely
and is gently curved. At the lower posterior end, which is little elevated above
the base line, the posterior margin passes in a stronger curve into the lower margin.
Lower margin very gently curved in its posterior part, sometimes almost straight
and subparallel to the upper margin, so that a lowest point cannot be located.
From about the middle it slopes upward, increasing the curve until it passes into
the anterior margin. Thus the anterior part of the shell appears only little narrower
than the posterior.

Valves very convex, convexity rather uniform over the disk, slightly stronger
over the posterior ridge, which is indistinct. The posterior slope is very slightly
compressed, without forming a wing-like expansion. Diameter 60 to 69 pr. ct.
of the length. Beaks much swollen and inflated, incurved, strongly elevated over

584

MEMOIRS OF THE CARNEGIE MUSEUM.

the lunula, and distinctly higher than the posterior upper margin, located at 28 to
33 pr. ct. of the length. Lunula cordiform, short and broad, almost as broad as
long.

Epidermis dull, with crowded, irregular, concentric lines. My specimens
being all dead shells, the original structure is somewhat doubtful, but in some speci-
mens there are distinct indications of a lamellar, cloth-like structure. No radial
sculpture present, and no distinct radial rib upon the posterior slope. Color of
epidermis paler or darker brown, rather uniform, often with a more or less distinct
blackish ra}^ upon the posterior slope, sometimes accompanied by a second ray.

Hinge-line gently curved or almost straight behind the beaks. Under the
lieaks it is sinuated, curving down more or less strongly, and then it is straight or
may curve up a little, rendering the lunula slightly concave. Pseudocardinal
teeth of the usual type, but somewhat variable; that of the right valve in particular
may be more or less compressed. Ligamental sinus rather small, triangular, its
anterior margin oblique.

Nacre white, shining in well preserved shells, iridescent, and with rather dis-
tinct radial lines. Prismatic zone narrow, without any trace of a sudden widening.
Cavity of shell and beaks considerable. Anterior adductor-scar deeply impressed.
Anterior retractor-scar small, close to the adductor-scar, but distinctly separated
from it. Anterior protractor-scar united with adductor-scar. Posterior adductor-
scar faint, siibovate, with a triangular upper process formed by the posterior
retractor-scar. No dorsal scars. Pallial line subconcentric to the margin.

JMeasurements.

No.

Length.

Height.

Diamoter.

Beaks

Figured.

1

22 nun.

18 Him.

= 82 pr. ct. of L.

14 mm. =64 pr. ct. of L.

at 6.5

mm. =30 pr. ct. of L.

PI. XL, fig. 6.

5

28 “

24 “

= 86

19 “ =68

8

“ =29

PI. XL, fig. 5.

7. . . .

30 “

25 “

= 83

18 “ =60

9

“ =30

10

41 "

33 “

= 80

27 “ =66

13.5

“ =33

PI. XL, fig. 4.

11

4!) “

30 “

= 80

34 “ =69

14

“ =29

12

57 “

43 “

=75

35 “ =61

16

“ =28

Remarks. — There cannot be any mistake about this species, which differs from
all others by the greatly swollen valves and inflated beaks, which render the shape
almost subglobular. The diameter of 60 to 69 pr. ct. is not found in any other
species of the genus, and approached only liy M . paraguayana, which, however,
distinctly differs in shape, being subangular, strongly oblique, and having the
posterior slope much comiiressed and subalate. In addition, the present species
differs from others by the narrow iirismatic zone. It is impossible to give exact
figures for the location of the greatest height, as the height of the shell is essentially
the same for a considerable disfance behind the middle.

ortmann: south American naiades.

585

Only one of the described species conies near to the present one, and this is
the little known M. inermis (Spix), as re-described by Von Ihering (1890, p. 126,
PL 9, figs. 1-3). This is founded upon a single, and apparently very young, in-
dividual, which resembles to a degree our youngest specimens of obesa. Von Ihering
gives the following figures for this:

Length. Height. Diameter.

21 mm. 15 mm. = 71 pr. ct. of L. 10 mm. =48 pr. ct. of L.

This specimen is not so high, and much less swollen than obesa, and we could
not by any means place it at the head of our table of measurements.

47. MoNOCONDYLiEA HOLLANDi Ortmaim, sp. nov.

Shell: Plate XLI, fig. 1.

Type-locality. — Sand-bar of Rio Guapore, near Rio Sao Simao, Matto Grosso,
Brazil. (J. D. Haseman coll., July 20, 1909). Type: Cam. Mus., Cat. No. 61.
5846. One specimen, male, with soft parts.

Characters of the Shell. — Shell large (length 102 mm.) moderately thick, out-
line subcircular, almost subrhomboidal. Height 77 pr. ct. of length. Valves
very little gaping, almost closed in front. Dorsal margin behind beaks practically
straight, in front of them subconcave, short and much lower, passing by a very
blunt angle into the anterior margin. The angle with the posterior margin is
also very blunt, almost regularly rounded. Posterior margin obliquely descending
and gently curved, passing in a sharper curve into the lower margin, thus forming
the rounded posterior end of the shell, which is situated a good deal above the base-
line. Lower margin curved, strongest curve in about the middle of the shell,
forming a blunt lowermost point at 47 pr. ct. of the length (consequently a little in
front of the middle). From this point the lower margin slopes iq) in either direc-
tion-; the posterior part of the margin is almost straight; the anterior more curved,
and passes in a regularly increasing curve into the anterior margin. The anterior
end of the shell cannot be called narrower than the posterior.

Valves gently and regularly convex in the middle of the disk, and more so
towards the beaks, distinctly compressed anteriorly and iiosteriorly, and the an-
terior compression is quite remarkable, forming a sharp, elevated, almost wing-
like expansion at the anterior upper margin. Posterior ridge of shell not at all
marked. Diameter 45 pr. ct. of length. Beaks somewhat inflated, not very
prominent (they are eroded), located at about 37 pr. ct. of the length. Lunula
short and broad.

Epidermis smooth, with irregular, concentric lines, poorly developed in the

586

MEMOIRS OF THE CARNEGIE MUSEUM.

middle of the disk and toward the beaks, slightly lamellar and more crowded on
the posterior slope and near the lower margin. Uiion the main part of the disk
there are traces of indistinct radial lines. Posterior slope with an indistinct radial
ridge. Color of epidermis yellowish brown, lighter in anterior half of the shell,
darker in the jiosterior, the two colors divided by a rather distinct radial boundary-
line through the middle of the shell. No color-rays are visible.

Hinge-line straight l:»ehind the beaks. In front it is curved down, and then
it slopes up again, making the lunula slightly concave. One pseudocardinal tooth
in each valve, that of the left valve standing in front of, and slightly below, that
of the right. The former is vertically depressed and projects into a slight groove
of the right valve. This groove has a low horizontal ridge, fitting into a groove
at the l^ase of the upper face of the tooth of the left valve; however, this may be
an individual character. Tooth of the right valve subpyramidal, not depressed,
projecting and fitting into a groove which is partly under the margin of the left
valve (the stumpy character of this tooth may also be individual). Ligamental

sinus triangular, about as deep as wide, its anterior margin oblique to the hinge-
line.

Nacre white, not very iridescent,"® with hardly any traces of radial lines.
Prismatic zone coinjiaratively narrow, slightly wider in the anterior part of the
shell, but not abruptly widening anywhere.

Cavity of shell shallow, that of beaks moderate. Anterior adductor-scar
sharply marked, elliiitical. Anterior retractor-scar above it and close to it, but
separated; in the right valve it ajipears double. Anterior protractor-scar con-
nected with adductor-scar. Posterior adductor-scar less sharply marked, subovate,
with a triangular jirocess aliove formed by the posterior retractor-scar. No dorsal
scars. Pallial line subconcentric to margin.

Measurements.

X

CJ

cc

I..ength.

Height.

Diameter.

Beaks.

Greatest Heiglit.

Figured.

d'

102 inni.

79 mm. =77 pr. ct. of L.

40 mm. =45 pr. ct. of L. at 38 mm. =37 pr. ct. of L. at 48 mm. =47 pr. ct. of L. PI. XLI ,
i 1 1 fig. 1.

Reiliarks. — A very striking and certainly new species, although only a single
individual is at hand. It is much larger than any of the known species, and has
a shape characterized chiefly by the location of the greatest height in front of the
middle, and the absence of a distinct narrowing of the anterior part of the shell.

The nacre ap])areiitilj'^ is slightly corroded. Although the specimen was alive when found, and
preserved with the soft parts, the up])ermost layer of the nacre has been injured and is exfoliating. Ac-
cording to my experience, this happens when specimens are allowed to die before they are put in alcohol.

ortmann: south American naiades.

587

It is also remarkable for the yellowish brown color of the epidermis, but this may
be variable. In the relative dimensions it comes near M. lentiformis (height and
diameter), and it is also a decidedly flat shell, although the beaks are slightly more
inflated than in lentiformis. The compression of the anterior extremity also should
be noted.

I name this species in honor of Dr. W. J. Holland, Director of the Carnegie
Museum, under whose auspices the expedition to Central South America by Mr.
John D. Haseman was made.

Anatomy. — The specimen at hand is a male.

Soft parts like those of the foregoing species, but the following points should
be mentioned : the inner edge of the anal opening is practically smooth, that of the
branchial opening with very minute papillae, which ajipear as mere crcnulations;
the gills have solid septa, which are unequal in thickness, heavier and lighter
ones alternating in a more or less regular way, chiefly so in the middle of the gills.

Genus Anodontites Bruguiere (1792).

Anodontites Bruguiere, Journ. Hist. Nat., Paris I, 1792, p. 131; Pilsbry, 1911, p.

609; Ortmann, 1911c, p. 91; Simpson, 1914, p. 1403.

Fatularia Swainson, Malacology, 1840, p. 287, 381.

Glabaris Gray (1847) Simpson, 1900, p. 916.

In this genus the hinge is without any teeth. From Mycetopoda and Leila it
differs by the absence of the characters peculiar to these, i.e., in the shape of the
shell, and certain features of the soft parts (See key p. 568). Furthermore we
find in Anodontites an extreme variability in the shape of the shell, from rounded
and subovate, to subtrapezoidal and elongated. The number of species is very
great, and it is hard to arrange them. Simpson (1914) distinguishes three sections.

1. Section Anodontites (sensu stricto). Shell rounded to elliptical; posterior
ridge low or wanting.

2. Section Styganodon Von Martens (1900). Shell subrhomboidal, with a
thick, dark, rather rough, sombre-colored epidermis, which is sometimes faintly
rayed; nacre lurid, shaded green.

3. Section Virguta Simpson (1900). Shell subsolid to solid, moderately in-
flated, greatly elongated, straight or falcate, rounded in front, sharply pointed
at the posterior base, where the high, sharply defined posterior ridge ends, and
above which it is somewhat obliquel^y truncated; beaks not high; epidermis green
to olive; nacre brilliant, blueish or purplish, iridescent, rayed with very fine, in-
distinct ridges; posterior end with a slight sinus.

588

MEMOIRS OF THE CARNEGIE MUSEUM.

It is seen at a glance that these three sections are not uniforml}- well defined.
Virgula, indeed, is sharply separated from the rest. Stygmiodon is well character-
ized by the epidermis; but unfortunately the type of the genus (Anondontites crispata)
undoubtedly belongs to Stygmiodon, having an epidermis (thick, dark, rough,
sombre-colored) which represents an extreme development of the Styganodon-
structure; in other characters also, A. crispata is closely allied to A. tenebricosa,“Hh.e
type of Styganodon.

It is clear that, on the one hand, Anodontites (sensu strictiore) must be used for
crispata, and, on the other hand, that Styganodon is a synonym of this, the type
of the latter being closely related to crispata. This necessitates a re-arrangement
of the sections, and a revision of their nomenclature. Although I have good
material representing the genus, it is impossible for me to attempt a final classifi-
cation, and the one given below is primarily adapted to the material at hand. An
attempt is made to preserve Simpson’s groups as far as possible.

Geographical distribution: The genus is widely distributed over South America
east of the Cordilleras from Patagonia to the Caribbean Sea, being found also in the
northern parts, where Diplodon and the Hyriinee in general are rare, or absent.
West of the Cordilleras it is generally missing, but it has been reported from that
side in Ecuador (the fact, however, requires confirmation). In addition the genus
has extended its range northward into Central America and Mexico, where it
is found in both the Pacific and Atlantic drainages.

Key to the Groups of Anodontites.

«!. SheJl not greatly elongated, nor pointed behind, without sharp posterior ridge. Posterior retractor-

scar connected with adductor-scar Subgenus Anodontites s. s.

bi. Epidermis dull, densely WTinkled.

Cl. Shell more or less elongated, but not distinctly oblique and subcircular.

di. Shell subtrapezoidal, somewhat elongated. Lower margin straight or concave.

Group of A. crispata.

di. Shell subovate or subelliptical, rather short. Lower margin convex.

Group of A. obtusa.

Cl. Shell distinctly oblique, short and high, nearly subcircular Group of A. trapezea.

bi. Epidermis more or less shining, wrinkles only partially developed or absent. Shell straight or
oblique, often distinctly so.

Cl. Shell not very oblicpie; straight, rather elongated, more or less pointed behind. Prismatic
border narrow, of eipial width Group of *4. trigona.

Cl. Sliell strongly oblique, not much elongated, not, or very little, pointed behind.
di. Shell subovate or subrotund. Prismatic border wide, width unequal.

Group of A. patagonica.

2’’ Another species, A. napoensis, allied to crispata, has been placed in Styganodon by Haas (1916,
pp. 32, 55).

ortmann: south American naiades.

589

di. Shell subovate or subtrapezoidal. Prismatic border narrow and of equal width.

Group of A. irapesialis.

ai. Shell greatly elongated, sharply pointed behind, with a sharp posterior ridge. Posterior retractor-
scar comidetely separated from adductor-scar. Epidermis not shining, covered with fine wrinkles.

Subgenus Lamproscapha.

Subgenus Anodontites s. s.

Aiiodontites s. s. + Styganodon Von Martens, Simpson, 1914, pp. 1403, 1448.

Shell of various shapes, but not greatly elongated or pointed behind, without
sharp posterior ridge. Posterior retractor-scar connected with adductor-scar.

1. Group of Anodontites crispata.

Group of A. crispatus Simpson, 1914, p. 1414 {pars) -f Section Styga7iodon, group
of A. tcnehricosus, Simpson, ibid., p. 1448.

Shell not, or very little, oblique, subtrapezoidal, rather elongated, with the
lower margin straight or more or less concave (sinuated). Epidermis dull, dark,
not rayed, not smooth, but strongly and densely sculptured all over by concentric
or radial, or irregular wrinkles. Prismatic border narrow or wider, of nearly equal
width.

The most essential feature of this group is in the texture of the epidermis,
and its somber color. The nacre also is peculiar, being dull and lurid.

These forms greatly resemble certain African species of Spatha, subgenus
Aspatharia, and I am strongly inclined to think that of all South American types
of Mutelmm these show the closest affinities to that African group. In A. crispata
occasionally a single dorsal muscle-scar is present, which also agrees with the con-
dition regularly seen in Spatha.

Three species belonging here are known to me in nature. They may be dis-
tinguished as follows :

ai. Wrinkles of epidermis arranged concentrically and radially, the railial tendency prevailing. Pris-

matic border moderate A. crispata.

a2. Wrinkles of epidermis with concentric tendency prevailing, radial arrangement obscure.

6i. Shell comparatively shorter and higher, beaks more median in position A. temhricosa.

ho. Shell comparative!}^ longer and lower, beaks more anterior in position A. clessini.

48. Anondontites crispata^*^ Bruguiere (1792).

Shells: Plate XL, figs. 7, 8; Plate XLI, figs. 2, 3.

Anodontites crispata Bruguiere, Journ. Hist. Nat. Paris, I, 1792, p. 131; Simpson,
1914, p. 1415.

Anodon r.eticulatus Sowerby, XVll, 1867, PI. 10, fig. 27.

Bruguiere uses Anodontites as feminini generis, and this should not be changed.

590

MEMOIRS OP THE CARNEGIE MUSEUM.

Glabaris crispatiis Simpson, 1900, p. 919.

Type-locality. — South America.

Other Localities. — Cayenne (Lea, Syn., 1870, p. 106); Amazon River (Sowerby,
reticulatus) .

New Locality. — Rio de la Paila, Paila, U. S. of Colombia (C. H. Eigenmann
coll., 1912). About twenty specimens, ten of them with soft parts, males and
females. (The Rio de la Paila is a tributary of the upper Rio Cauca of the Rio
]\lagdalena-drainage) .

Distribution . — Simpson says that this species is widely distributed in tropical
South America, though but few exact localities are known.

Characters of the Shell. — Of medium size (maximum length 67 mm.), moder-
ately thick, rather thin when young. Outline elongated subtrapezoidal, or sub-
elliptical. Height 52 to 61 pr. ct. of the length. Valves practically closed, or very
little gaping in front. Dorsal margin straight, or prevalently very gently curved
behind the beaks, descending in front of the beaks, and passing gradually into
the anterior margin. Posteriorly it forms a blunt angle with the posterior margin,
or passes into it in a curve. Posterior margin curved, obliquely descending in
its upper part, becoming gradually steeper, and sometimes nearly vertical at the
posterior end, which is blunt and rounded, but bends rather suddenly into the
lower margin. The posterior end is thus very little elevated above the base-line,
often practically at its level. Lower margin straight for a considerable distance,
often even slightly concave in the middle, gently sloping upward toward the front
and curving up into the anterior margin. This ascending part, and the descending
anterior upper margin, make the anterior end of the shell narrower, but the whole
shell does not appear very obli(|ue. The highest part of the shell is in the posterior
section.

Valves moderately convex, rather flattened upon the sides, and sometimes
even with a shallow depression corresponding to the emargination of the lower
margin. Posterior ridge broad and rounded, indistinct, but the greatest diameter
of the shell (31 to 38 pr. ct. of length) is situated upon it, so that the shell is more
swollen posteriorly than anteriorly. Posterior slope slightly compressed, sometimes
with a very faint radial groove. Beaks not swollen and not elevated above the
hinge-line, located at 27 to 33 pr. ct. of the length. Lunula indistinct or narrow,
not very long.

Epidermis finely wrinkled all over, the wrinkles partly concentric, but pre-
vailingl\' arranged in a radial pattern. Concentric and lamellar wrinkles are found
chiefly near the margins. Upon the disk, the pattern varies in the anterior and

ortmann: south American naiades.

591

posterior section of the shell. Anteriorly there are short, concentric wrinkles,
arranged in scalariform, radial bands. Posteriorly the wrinkles are rather irregu-
larly radial, assuming generally an oblique direction, forming radial liands of
loops or V-shaped festoons, often anastomosing, and more or less reticular (“like
dried paint”). In the middle of the shell the two types of sculpture pass into each
other: the short, subconcentric, scalariform wrinkles become rather suddenly V-
shaped, and assume the oblique or radial direction. On the posterior slope and
near the lower margin, the wrinkled sculpture is more or less obscured by sub-
lamellar concentric striae. Color of epidermis, in very young specimens, yellowish;
later it becomes brownish olive or dark greenish to blackish, generally more brown-
ish toward the beaks, and more blackish toward the margins. Sometimes (in
strong transmitted light) there are dark color raj^s upon the iiosterior slope.

Hinge-line nearly straight or gently convex posteriorly, descending anteriorly,
a little irregular just in front of the beaks. Ligamental sinus moderate, triangular,
wider than deep, its anterior margin slightly oblique to the hinge-line. Nacre
blueish white, grayish white, lurid, often showing brownish or greenish discoloration.
Prismatic border rather narrow, subequal in width, of grayish color.

Cavity of shell and beaks shallow. Anterior adductor-scar well marked, im-
pressed, irregular!}^ elliptical. Anterior retractor-scar only partially separated
from adductor-scar, connected with it narrowly or more broadly, and very variable
in this respect. Anterior protractor-scar separated from adductor-scar. Pos-
terior adductor-scar less imiiressed, rounded or subovate, posterior retractor-
scar forming an upper jirocess of it. Dorsal scars generally absent, but in a few
cases there is an indistinct single one. Pallial line tlistinct, simjile, subparallel
to the margin.

Measurements.

No.

Sex.

Length.

Height.

Di.ameter.

Beaks.

Figured.

1

d'

24.5

imii.

15 mm. =61 pr. ct. of L.

8.5

mm. =35 pr. ct. of L.

at 7 mm. =29 pr.

•.t. of L.

37

19.5 “ =53

11.5

“ =31

10 “ =27

PI. XLI, fig. 3.

2

9

45

27.5 “ =61

17

“ =38

15 “ =33

5

9

5G

29 “ =52

19.5

“ =35

15 “ =27

10

9

63.5

35 “ =55 “

22

“ =35

IS “ =2S

PI. XL, fig. 8.

?

67

35 “ =52

22

“ =33

19 “ =28

PI. XLI, fig. 2.

Simpson

53

27.5 “ =52

17

“ =32

Reynarks. — I have no doubt that my specimens represent A. crispata. They
agree fairly well with Sowerby’s figure of Anodon reticulatus, and very well with
Simpson’s description. But there are several closely allied, if not identical, species,
chiefly A. napoensis Lea (Obs. XII, 1809, PL 53, fig. 137, and Germain, 1910, p.
C 64, PI. 2, figs. 3, 4) from Rio Napo (tributary to the upper Amazon in Ecuador)
and Rio Unuyacu (tributary to Napo). This s]iecies differs chiefly in the posterior

592

MEMOIRS OP THE CARNEGIE MUSEUM.

end of the shell, which is more elevated above the base-line, and in the more curved
ventral margin. The sculpture of this species undoubtedly is similar, but the
details have not been described.

Sim})son says that this species is very variable in sculpture. However, I
find that my s])ecimens (twenty at hand) are rather uniform in this respect. Only
in the youngest specimen is the subradial sciiljiture of the posterior part not so
well developed; but in the second (37 mm. long) it is distinctly seen.

There is a good deal of variation in the shape of the posterior end, and the
liosterior margin may be more oblique or may be more vertical in its lower part.
Very often the nearly vertical truncation of the jiosterior end produces the ap-
pearance of a biangulation, chiefly when the faint groove of the posterior slope is
visible.

Anatomy. — Soft parts of six males and four barren females at hand for study.
Anal opening sejiarated from the branchial by a mantle connection, open and
nowhere closed, its inner edge smooth or nearly so. Branchial opening with fine
jiapillae. Paljii longer than high, lower margins convex, posteriorly truncated,
the truncation forming the short jiosterior margins, which arc connected at base
only.

Gills of medium width, the inner the wider anteriorly, its anterior end im-
mediately behind the jialpi, and attached to the whole interval between the palpi
and the anterior end of the outer gill. The latter at the highest ])oint of the mantle-
attachment-line. Inner lamina of inner gill entirely connected with abdominal
sac.

Gills with well develoiied se])ta. In the non-marsupial gills the septa are
irregularly alternating, stronger and weaker. In the female the inner gill is mar-
supial for nearly its whole length, with stronger and more uniform, but not more
crowded, sejita. The septa possess the usual swelling, forming vertical ridges
projecting into the lumen of the water-tubes, dividing the latter into an inner
compartment (ovisac), and an outer (secondary water-tube).

49. Anodontites tenebricosa (Lea) (1834).

Anodonta tenebricosa. Lea, Obs., I, 1834, PI. 12, fig. 36; D’Orbigny, 1843, p. 616.
Anodon te7iehrosa Sowerby, XVII, 1867, PI. 13, fig. 43.

A'tiodon tenebricosa Bgwerby, XVII, 1870, PL 31, fig. 123.

Anodonta tenebricosta Corsi, 1901, p. 457, fig. 58.

Glabaris tenebricosa Von Ihering, 1893, p. 61; Nehring, 1893, p. 163; Simpson^

1900, p. 930.

ortmann: south American naiades.

593

Anodontites tenebricosus Simpson, 1914, p. 1448.

Anodontites (Styganodon) tenebricosus Haas, 1916, pp. 35, 57.

Type-locality. — Rio Parana.

Other Localities. — Rio San Jose and Arroyo del Rosario, Uruguay (D’Orbigny)
(tributaries to La Plata in the Banda Oriental); Rio de la Plata, Buenos Aires,
Argentina (D’Orbigny); Buenos Aires (Haas); Rio Uruguay, Salto Oriental, Uru-
guay (Haas); Rio Piracicaba, Piracicaba, Sao Paulo, Brazil (Von Ihering) (Nehring).

New Localities. — Pond along banks of Rio Negro, Santa Isabel, Uruguay (J.
D. Haseman colL, February 11, 1909). One complete specimen and one left valve.
Rio de la Plata, San Isidro, 20 km. North of Buenos Aires, Argentina (A. Wind-
hausen coll., January, 1917). One female with soft parts.

Distribution. — From the region of Buenos Aires in the La Plata system and
its tributaries in Uruguay (smaller streams of the Banda Oriental and Rio Uruguay-
system) up the Parana River to its headwaters in Sao Paulo, Brazil.

Simpson gives a much larger range, adding even Ecuador and Peru, but I do
not know on what authority.

This species is well known and has been well described. Its chief characters
are its shajie, which, although elongated-subtrapczoidal, is rather short and high
in comparison with the related species. The prismatic border of the shell, chiefly
in old shells, is remarkably wide, but subequal in width, not suddenly changing
an}' where.

The character of the epidermis has been given as concentrically lamellose.
This is correct, and the fine, crowded, concentric lamellae are very obvious. How-
ever, closer investigation shows that between these lamellae are additional, more
irregular wrinkles, which produce upon the posterior part of the shell a reticulated
(anastomosing) sculpture, and more anteriorly these wrinkles may be even radial.
But they exist only between the concentric lamellae, and thus they are very short
and largely obscured by the concentric scul]3ture. This difference of sculpture
from that of A. crispata is very striking. Otherwise these two species have many
features in common.

Measurements.

1 Sex.

Length.

Height.

Diameter.

Beaks.

S. Isabel

S. Isidro

!

i 9

57 mm.
92 “

33 mm. =58 pr. ot. of L.
54 “ =59

17 mm. =29 pr. ct. of L.
32 “ =35

at 18 mm. =32 pr. ct. of L.
27 “ =29

According to Von Ihering’s measurements, the height is 53 pr. ct. and the
diameter 34 pr. ct. of length. According to Nehring’s measurements the height is
48 to 55 pr. ct., the diameter 30 to 33 pr. ct. Thus specimens from the upper

594

MEMOIRS OP THE CARNEGIE MUSEUM.

Parana-drainage seem to be not so high, more elongated than those from the lower
part of the system. In Lea’s original specimen (according to figure and measure-
ments given by Simpson), the height is 55 pr. ct., the diameter 45 ]ir. ct. of length.
In another specimen measured by Simpson, the height is GO pr. ct., the diameter
38 ])r. ct. D’Orliigny’s measurements give for height 58 pr. ct. and for diameter
37 ])r. ct. These latter figures agree very well with mine, and it should be noted
that the larger specimens generally give a proportionally greater diameter.

Anatomy. — The soft parts of one barren female are at hand for examination.
Practically identical with the preceding species. The gills, in the present
specimen, ajiiiear comiiaratively narrow. The structure of the rnarsupiiim (inner
gill) is typical. The swellings of the sejita are very distinct, and arc situated closer
to the outer lamina (primary limb).

50. Anodontites clessini (Fischer) (1890).

Shells: Plate XLI, fig. 4; Plate XLII, figs. 1, 2. Anatomy of gills: Plate XLVII,

fig. 4.

Mycetopus plicatus Clessin, Malakozool. Blrett., IT, 1882, p. 190, PI. 4, fig. 7
{no7nen prwoccupatum) .

Mycetopus clessini Fischer, Journ. de Conchylioh, XXXVIII, 1890, p. 8, footnote.
Glaharis nehringi Von Ihering, 1893, j). GO; Nehring, 1893, p. 1G3; Von Ihering,
1910, ]). 139.

Glaharis clessitii Simpson, 1900, p. 930.

A nodontites clessini Simpson, 1914, p. 1450.

Type-locality : ?

Other Localities. — Rio Sta. Maria, Rio Grande do Sul, Brazil (Von Ihering)
(tributary to Ibicuhy and Uruguay); Rio Piracicaba, Piracicaba, Sao Paulo, Brazil
(Von Ihering); Rio Piracicaba Vlirim, Piracicaba, Sao Paulo, Brazil (Nehring);
Rio Paraguassu, Bahia, Brazil (Von Ihering, 1910).

New Locality. — Rio Vaccahy Mirim, Santa Maria (da Bocca do Monte),
Rio Grande do Sul, Brazil (J. D. Ilaseman coll., January 29, 1909). Twelve speci-
mens, all with soft jiarts, males and females.

Distrihulion. — Positively known from the Uruguay drainage and the head-
waters of the Parana. In addition, it has crossed over into certain coastal streams
in Brazil. Our locality in Rio Vaccahy Mirim belongs to the Jacuhy-Guahyba-
system, but is close to Von Ihcring’s record from the Uruguay drainage, but on
the other side of the divide. This species has been found also in the Rio Paraguassu
in Bahia, well to the North, and separated from the rest of the range in the upper
Parana (Piracicalia). But ]irobably some kind of connection will be found.

ortmann: south American naiades.

595

This species may be regarded as an elongated and narrow tenebricosa. In
consequence of the general elongation/the posterior margin forms a more obtuse
angle with the upper margin, and the posterior end is not sulitruncated, but more
evenly and narrowly rounded. Young specimens are almost regularly long-
elliptical. The prismatic zone is comparatively narrow. The ligamental sinus is
triangular, wider than deep, and its anterior margin forms an obtuse angle with
the hinge-line (being directed obliquely backwards), or is almost vertical. (In
A. tenebricosa this sinus is about as dee]) as wide, its anterioi- margin is nearly
vertical and curved gently forwards, so that the lower point, which is (]uite sharp,
is directed obliquely forward).

In all other characters the two s])ecies resemble each other, and this is pre-
eminentl}^ true of the sculpture of the e])idermis.

IMeasurements.

No.

Sex.

Length.

Height.

Diametei .

Beaks.

Figured.

1. .

cf

31 mm.

16 mm. =52 pr. ct. of L.

8.5 mm. =24 pr. ct. of L.

at 10 mm. =32 pr. ct. of L.

4. .

c?

45 “

22 “ =49

13 “ =29

12 “ =26

8. .

9

Cl “

27 “ =44

19 “ =31

17 “ =28

11. .

9

68 “

34 “ =50

20 “ =29

19 “ =28

PI. XLII, fig. 2.

10. .

9

C9 “

31 ‘‘ =45

23 “ =33

18 " =26

PI. XLl, fig. 4;

12. .

&

70 “

32 “ =46

21 “ =30

18 “ =26

PI. XLII, fig. 1.

According to Von Ihering, the height is from 43 to 51 pr. ct., the diameter from
22 to 33 pr. ct. of length, the beaks are at 24 to 31 pr. ct. According to Nehring,
the height is 41 to 43 pr. ct., the diameter 20 to 25 ])r. ct. In A. tenebricosa, the
figures for the height are 55 to GO i)r. ct. (rarely below this); for the diameter 29
to 45 pr. ct. and for the beaks 29 to 34 pr. ct.

Remarks. — The greater length of the shell is brought out liy these figures,
and also the lesser obesity and more anterior position of the beaks, which results
from it. It is also seen in my specimens that the elongation is not so great in
younger specimens, and in the latter the measurements approach those of tenebricosa,
or even fall within the range of variation of it. It is possible that the two species
actually intergrade, a condition which has been hinted at by Nehring (p. 1G4) to
exist in the Rio Piracicaba.

Anatomy. — The soft jiarts of seven males and five barren females have been
investigated.

Color of- soft parts whitish; inner edge of anal and branchial openings black,
the black color running forwards from the branchial for a certain distance.

Anal opening entirely oiien, its inner edge practically smooth, sejiarated
from the branchial opening liy a connection of the mantle-margins. Inner edge

596

MEMOIRS OF THE CARNEGIE MUSEUM.

of branchial opening with distinct, but small papillae. Palpi rather small, semi-
circular, shortly truncated iiosteriorly, thus forming posterior margins, which are
not connected.

Gills (PI. XLVII, figs. 4a, h) long and narrow, the inner considerably wider
than the outer, chiefly anteriorly. Anterior end of inner gill immediately behind
the palpi. Inner lamina of inner gill entirely connected with abdominal sac.
Septa well develo]ied, those of the non-marsupial gills alternately somewhat stronger
and weaker, best seen in the outer gill of the female (PI. XLVII, fig. 46). Inner
gill of female marsupial for nearly its whole length, with more uniform, thicker,
but not more crowded septa, which have the usual swellings near their contact
with the outer lamina.

2. Group of Anodontites ohtusa.

Simpson, 1914, p. 1453.

Shell not very oblique, subovate or sulDelliptical, rather short, lower margin
convex. Ejiidermis dull, greenish, often with rays, concentrically lamellarly
wrinkled. Prismatic liorder narrow.

The few forms belonging here are closely allied to the first group, and have
indeed been ])laced in the section of Styganodon by Simpson. They are shorter,
higher than those of the crfspa/a-group, with more inflated beaks, and have an
unusual development of color-rays. The sculptui-e of the epidermis is much like
that of tenehricosa. and clessini, and consists of close, sublamellar, concentric wrinkles,
here and there subreticulated.

51. Anodontites obtusa (Spix) (1827).

Anodon obtusum Spin, 1827, PI. 22, fig. 3; Wagner, Ibid, p. 30.

Anodo7i lituraium Spin, Ibid, PI. 22, fig. 4.

Anodonta ohtusa and litturata Hupe, 1857, pp. 86, 87, PI. 17, fig. 4.

Anodon ohtiisus Sowerby, XVII, 1867, PI. 12, fig. 39.

Anodcm liturata Sowerby, Ibid, 1868, PL 20, fig. 78.

Anodonta ohtusa Von Ihering, 1890, ji. 159.

Glaharis ohtusiis and lituratus Simpson, 1900, p. 931.

Anodonta {Gluharis) ohtusa Germain, 1910, p. 63, PL 3, figs. 14, 15.

Anodontites ohtusus and lituratus Simpson, 1914, pp. 1453, 1454.

Type-locality . — Rio Paraguassu, Bahia, Brazil.

Other Records. — Rio Paraguassu (Von Ihering, 1910, p. 139); Rio Sao Francisco,
Villa Nova, Sergijie, Brazil (Von Ihering); Rio Sao Francisco, Joazeiro, Bahia,
Brazil (Von Ihering); Rio das Velhas, Minas Geraes, Brazil (Von Ihering) (upper

ortmann: south American naiades.

597

S. Francisco-drainage) ; Bodegas, and Rio Daiile, Ecuador (Germain) (Pacific-
drainage, near Guayaquil).

New Locality. — Lagoa Saclio Grande, Cidade da Barra, Bahia, Brazil (J. D.
Haseman coll., December 24, 1907). One young specimen. (S. Francisco-
drainage) .

Another, larger specimen is in the Carnegie Museum, from the Holland Collec-
tion, labeled “Brazil.”

Distribution. — Known from the drainages of Rio Paraguassu and Sao Fran-
cisco. Von Ihering gives a possible variety, similar to the var. hohenackeri, from
Rio Mucury, Bahia (southern part) (1890, p. 161), and another variety {juparana,
1910, p. 131) from Lagoa Juparana of Rio Doce, Espirito Santo). The form
is not known from the basin of the Amazon. So much more astonishing is Germain’s
record of this species from the Pacific slope in Ecuador. I cannot discover any
differences in these specimens, but attention should be called in this connection
to A. aff. pastasanus of Haas (1916, pp. 34, 56, PI. 2, fig. 1) which comes from the
identical localities (Rio Daule and Bodegas in Ecuador). The latter, however,
is more elongated than Germain’s figure of obtusa, and cannot be the same.

Simpson also cites Paraguay, but I do not know on what authority.

After what Wagner and Von Ihering have said, it is perfectly clear that liturata
is the young stage of obtusa, and my larger specimen shows, near the beaks, the
juvenile character of broken and oblique rays. Hupe says that liturata is less
swollen than obtusa, while Simpson (1914, p. 1454) states the opposite, that it is
more inflated (possibly slip of the pen). Of my two specimens it is the smaller
one, with ZR^rata-color-markings, which is the more swollen.

Measurements.

Length.

Height.

Diameter.

Beaks.

Barra

Brazil

Spix’ type . .

36 mm.

47 “

53 “

24 mm. = 67 pr.
32 “ =68

35 “ =66

ct. of L.

ii

H

17 mm. = 47 pr. ct. of L.
19 “ =40

25 “ =47

at 12 mm. = 33 pr. ct. of L.
17 “ =36

3. Group of Anodontites trapezea.

Shell subcircular, but distinctly oblique, short and high. Epidermis dull,
densely, concentrically, lamellarly wrinkled. Prismatic border moderate, or narrow,
of nearly equal width.

The subcircular shape and the dull, cloth-like epidermis are found in no other
group combined.

In 1910 (p. 131) he says that specimens from Rio Mucury &vq juparana.

598

MEMOIRS OF THE CARNEGIE MUSEUM.

52. Anodontites TRAPEZEA (Spix) (1827).

Anodon rotundum and trapezemn Spix and Wagner, 1827, p. 28, PI. 20, figs. 1-4.
Anodonta rotimda and trapezea Von Ihering, 1890, pp. 142, 145, PI. 9, figs. 5, 6.
Anodonta cailliaudi Lea, Obs., X, 1863, PI. 45, fig. 297.

Anodon cailliaudi Sowerby, XVII, 1867, PI. 12; fig. 38.

Glabaris rotunda Von Ihering, 1893, p. 59; Simpson, 1900, p. 918.

Glabaris trapezea Von Ihering, 1893, p. 57; Nehring, 1893, p. 163; Von Ihering,

1910, p. 138.

Anodontites rotundas Simpson, 1914, p. 1410.

D’Orbigny first recognized the identity of trapezea and rotunda, and selected
the first name. Although the specimens from Corrientes, to which he applied
this name, may not have been the typical trapezea, but a variety or even a species
(spixi D’Orbign^y 1835), his selection of trapezea in preference to rotunda should
be accepted.

Type-loccdity . — Rio Solimoes (middle Amazon) Brazil.

Other Localities. — Rio vSao Francisco, Villa Nova, Sergipe, Brazil (Von Ihering,
trapezea)-, Sao Paulo, Brazil (Wagner, rotunda); Rio Piracicaba, Piracicaba, Sao
Paulo, Brazil (Von Ihering, trapezea).

This species, partly as rotunda, partly as trapezea, has been reported (D’Or-
bigny, Lea, Von Ihering) from the lower Rio Parana (near Corrientes). However,
the A. trapezea of D’Orliigny is believed to be a different form or variety {A. spixi
D’Orbigny). It is surely allied to trapezea.

New Localities. — Lagoa de Sacho Grande, Cidade da Barra, Bahia, Brazil
(J. D. Haseman coll., December 24, 1907). Six complete shells and four odd (left)
valves. Lagoa de Sacho Pecpieno, Cidade da Barra, Bahia, Brazil (J. D. Haseman
coll., December 24, 1907). Three specimens, one a male with soft parts. Rio
Grande, Ba'rreiras, Bahia, Brazil (J. D. Haseman coll., January 4, 1908). Three
males with soft parts. Rio Sao Francisco, Joazeiro, Bahia, Brazil (J. D. Haseman
coll., February 28, 1908). One odd (left) valve.

All these localities are in the drainage of the Rio Sao Francisco.

Distrihution. — The presence of this species in the system of Rio Sao Francisco
is established. Since Von Ihering has also cited it as trapezea from the upper
Parana (Piracicaba) in Sao Paulo, it must belong to both systems. Farther down
the Parana (Corrientes, Argentina) similar shells are jiresent, but apparently
slightly different from the type. The original locality is in the Amazon-drainage,
but it lias not subsecpiently been found there.

Gharacters of the Ahell. — Shell of moderate size (maximum length 75 mm.),

ortmann: south American naiades.

599

rather thin when young, slightly more solid when older, but never of consideralile
thickness. Outline subcircular, but a little irregular and variable, and distinctl}^
oblique. Height 80 to 93 pr. ct. of the length. Valves not gaping. Dorsal
margin nearly straight. Posterior and anterior margins uniting with the dorsal
in blunt angles. Posterior margin obliquely descending, first straight, then curved,
and passing in a rather regular curve in the posterior part of the lower margin,
without any trace of a posterior point. Lower margin ascending forward more
strongly, and curving up into the anterior margin, so that the anterior part of the
shell appears somewhat narrower than the posterior part, thus producing the
obliquity.

Valves inflated, diameter 49 to 56 pr. ct. of the length. Beaks somewhat
swollen and inflated, elevated above the hinge-line and incurved, their tips im-
mediately above the hinge-line in the young, a little higher in older shells. Loca-
tion of beaks at 33 to 44 pr. ct. of the length. Outer surface of shell rather regularly
convex, but anterior and posterior slopes somewhat compressed. Posterior ridge
quite indistinct.

This species is remarkable for the presence of beak-sculpture, which is generally
absent in this subfamily. However, I think that this sculpture is not genetically
connected with that of other Naiades, but probably is independently developed.
•Quite a number of my younger specimens show it. The very tip of the beak ap-
pears as a small tubercle, and is succeeded by three to five concentric bars, which
follow the growth-lines, and are low and rounded, but perfectly distinct in the
middle, disappearing anterior!}' and posteriorly. These bars are restricted to and
crowded together at the extremity of the beaks, and disappear at a short distance
(3 to 4 mm.) from them.

Epidermis with fine and crowded, concentric, somewhat anastomosing lines,
which become lamellarly elevated toward the margins, and in well jireserved shells,
chiefly young ones, they have this character all over the shell. In addition there
may be fine and faint radial strias, but there are no scalariform stripes. In old
and partly worn shells the surface becomes rather smooth, but remains always dull,
and is not shining. Color of epidermis from dark green to yellowish brown. The
normal color in young specimens seems to be lighter or darker green, sometimes
with indistinct dark green rays (seen only in transmitted light). In older specimens
the color becomes greenish brown to light brown, due to partial abrasion of the
epidermis. There are always two more or less distinct dark green or blackish
rays upon the posterior slope, often accompanied by two lighter, yellowish rays.
Larger specimens may have a few dark brown growth-rests.

600

MEMOIRS OF THE CARNEGIE MUSEUM.

Hinge-line practically straight in young shells; in older shells it curves gently
down under the beaks, and up again at the anterior end, and the posterior end
curves gently down, thus forming a slight S-curve. Ligamental sinus triangular,
not deeper than wide, varying with age (shallower in young shells), its anterior
margin running obliquely backward in the young, and vertically in older individuals;
its lower point may be sometimes directed forwards.

Cavity of shell and of the beaks rather deep, corresponding to the obesity
of the shell. Nacre white; in young shells blueish white, in older shells somewhat
inclining to cream-color, always extremely glossy, silvery, and iridescent toward the
margins, with fine, straight, and irregular radiating lines. Along tlie margin there
is a rather narrow nacreless (iirismatic) zone, relatively wider in young specimens.
This zone is subconcentric with the shell-margin, widest in the middle, gradually
narrowing towards the ends, but nowhere suddenly or markedly changing its width.

Anterior adductor-scar and anterior retractor-scar united, not deep, irregularly
ovate or elliptical; anterior protractor-scar connected with adductor-scar, or,
in old specimens, more or less (sometimes distinctly) separated. Posterior ad-
ductor-scar faint, subovate, the posterior retractor-scar forming an upper pro-
jection thereof. No dorsal scars. Pallial line subconcentric to margin.

IMeasurements.

‘ Location.

No.

Sex.

Length.

Height.,

Diameter.

Beaks.

Saclio C! ramie ....

a

?

30 nun.

24 mm. =S0 pr.

:t. of L.

15

mm. =50 pr. cl. of L.

at 13

mm. =43 pr. cl. of L.

Saclio Pcciueno . . .

c?

41

34 “ =83

21

“ =51

18

“ =44

Barreiras

1

d’

47

42 “ =89

24

“ =51

18

“ =38

Do

3

54

47 “ =87

27

“ =50

20

“ =37

Saclio Grande ....

b

?

5S

48 “ =83

30

“ =52

24

“ =41

Do

c

V

68 “

63 “ =93

38

" =56

30

“ =44

Sacho Pequcno . . .

V

72.5 “

01.5 “ =85

38

“ =52

24

“ =33

trapezea, type ....

03

52 “ =84

31

“ =49

(Von Ihering)

rotunda, type

42

30 “ =86

22

“ =52

Do.

trapezea (Piracioaba) . . .

00

49 “ =82

33

“ =55

Do.

rotunda (Simpson)

75

64 “ =85

38

“ =51

The variety (or species )sptxi D’Orbigny from the lower Parana at Corrientes
grows larger, reaching the length of 81 mm. according to Von Ihering, and of 90
mm. according to D’Orbigny.

Reviarks. — This shell is easily recognized by the general shape and proportions,
l)y the rather narrow prismatic border, and by the dull, greenish color of the epi-
dermis. That trapezea and rotunda are only the old and the young stages of the
same species, is conclusively shown by our material, chiefly by the sets from Cidade
da Barra. The old specimens have the beaks a little elevated above the hinge-
line, and the hinge-line is gently curved; while the young specimens have the point

ortmann: south American naiades.

601

of the beaks immediately above the hinge-line, and have the latter straight. Speci-
mens of intermediate size intergrade in these characters.

Anato7ny. — The soft parts at hand are not in good condition. However, the
structure of the gills can be made out, and according to the alternation of stronger
and weaker septa, all four specimens at hand are males.

In other respects the usual structure of the genus is seen. Anal opening en-
tirely open, separated from the branchial by a mantle-connection. Branchial
opening with distinct, but small papillae. Palpi small, subcircular, with a short
truncation at the posterior end; the posterior margins not connected. Inner lamina
of inner gill entirely connected with abdominal sac.

4. Group of Ariodontites trigona.

Simpson, 1914, p. 1441.

Shell not very oblique, rather elongated, subelliptical, or subovate, narrowly
rounded, or somewhat pointed behind. Epidermis more or less shining, and not
uniformly and densely covered with wrinkles, although such are present here and
there. Prismatic border narrow, of nearly equal width.

This group is poorly defined. Its chief character is the rather elongated shell,
somewhat pointed behind, or narrowly rounded, and not distinctly oblique. The
comparatively smooth e])iderniis is another noticeable feature, but still there are
species, which have sublamellar, concentric striae, at least in parts of the shell.
The color of the epidermis is not so sombre and dull as in the iireceding groups.
The narrow prismatic border seems to be constant.

53. Anodontites trigona (Spix) (1827).

Anodon triyonum Spix & Wagner, 1827, p. 29. PI. 22, fig. 2.

Glabaris trigonus Simpson, 1900, p. 928.

Anodontites trigonus Simpson, 1914, p. 1441.

Anodon moretonianus Sowerby, XVll, 1867, PI. 9, fig. 20.

Doubtfully synonymous:

Anodon georginoe Griffith, 1834, p. 595 (index), PL 19, fig. 3.

A. moretonianus Sowerby undoubtedly is this species. Simpson (1914, p. 1431)
is mistaken in placing it with A. trapesialis. Sowerby ’s moretonianus is not A.
mortoniana of Lea.

A. georginoi Griffith, from “rivers of Paraguay,” also seems to be this species.
The figure given l.c. is shorter and higher, but the characteristic shape and the
radial ribs (although too much emphasized) well agree with it. It is also from a
region, where trigona is known to occur. For this species, Simpson (1900, p. 927 ;
1914, p. 1440) creates a separate group.

602

MEMOIRS OF THE CARNEGIE MUSEUM.

Simpson makes Anodonta castelnaudi Hiipe a synonym of trigona, but I do not
think that this is correct. A. castelnaudi lacks the chief characteristic features of
trigona: the pointed jiosterior end and the rib upon the posterior slope.

Type-locality. — Rivers of the “province Rio Negro.” There is now no such
lu’ovince in Brazil, from which country the collections of Spix came.

Other Localities. — Amazonas and Bolivia (Von Ihering, 1893, p. 120); Rio
Xingu, Para, Brazil (tributary to lower Amazon) (Von Ihering, 1910, p. 137);
Tributaries of Amazon in Bolivia (territory of tlie Chiquitos and Moxos) (D’Or-
bigny); Rio Estacamento, Peru (Haas, 1916); Rio Paraguay, San Luis de Caceres,
iMatto Grosso, Brazil (Von Ihering, 1915, p. 13) ; Rio Batel and Rio Parana, Corri-
entes, Argentina (D’Orbigny).

N'ew Localities. — Swam]) of Lambare, Asuncion, Paraguay (J. D. Haseman
coll., March 31, 1909). One right valve. Headwaters of Rio Paraguay, Santa
Rita, Matto Grosso, Brazil (J. D. Haseman coll., June 12, 1909). Two specimens,
male and female, with soft parts. Rio Limay, Patagonia, Argentina (W. Israel
donor). One specimen.

Distribution. — According to Von Ihering (1890) : “Everywhere in the Amazonas
region, but also in the La Plata up to Corrientes.” Simpson gives: Brazil, Ecuador,
Peru, Bolivia. The species undoubtedly has a wide distribution, both in the
Amazon and the Paraguay-Parana drainages, but is apparently missing in the upper
Parana. The new locality in Rio Limay in Patagonia (Rio Negro-drainage) con-
siderably extends the southward range.

Description of Shell. — Shell rather thick and solid. Outline elongated-ovate,
pointed behind, lower margin convex, forming a bluntly projecting angle about
its middle. Height 59 to 61 pr. ct. of length according to my specimens (in D’Or-
bigny’s the height is only 54 pr. ct.). Valves not gaping. Dorsal margin gently
curved, posterior part almost straight, anterior part descending. Posterior angle
of dorsal margin obtuse, but well marked. Anteriorly the dorsal margin forms
a very indistinct angle, or passes gradually into the anterior margin. Posterior
margin descending oblicpiely, and almost straight or very gently curved, passing
into the iiosterior part of the lower margin in a very sharp curve, which forms the
jiosterior point of the shell. This point is somewhat elevated above the base-
line, since the jiosterior part of the lower margin slopes upward. This part of
the lower margin is almost straight. The lowermost point of the lower margin
is a little behind the middle, and in front of it the lower margin changes its direc-
tion, running upward and forward, so that this lower point forms a blunt projection.
The ascending anterior portion of the lower margin is at first almost straight or

ortmann: south American naiades.

603

very gently curved, and then it curves up into the anterior margin, which is narrowly
rounded. The anterior part of the shell does not appear narrower than the pos-
terior. The greatest height of the shell is a little behind the middle, and anteriorly
it rather gradually becomes narrower, while posteriorly it tapers vei^y decidedly
to the posterior point. Thus there is also no marked obliquity in the shell. (In
the young specimen from Santa Rita the posterior taper is not so strong).

Valves moderately convex, diameter 34 to 36 iir. ct. of length. Beaks not
very prominent above hinge-line, located at 18 to 26 pr. ct. of the length. Con-
vexity of the valves greatest over the posterior ridge, least between this ridge and
the region of the lower angle of the lower margin. Posterior slope compressed,
but hardly any compression at anterior end. A distinct radial rib upon the pos-
terior slope, lying between two radial depressions; sometimes there is a trace of a
weak radial rib above it, but generally this is not very distinct. In young specimens
the radial rib is indistinct.

Epidermis rather smooth, with irregular concentric lines, which become
lamellar upon the posterior slojie and towards the ventral margin. No distinct
radial sculpture, except some irregular and rather fine scalariform stripes. Color
of epidermis normally a very dark green upon the disk, which, however, may turn
to brown. The posterior slope may also be dark green, with the shallow grooves
brownish, or it may be entirely brown, and in the young specimen it is brown,
with the radial rib marked b}^ a rather distinct dark green ray. The disk has no
color rays.

Hinge-line practically straight behind the beaks, and in front of them it
gently curves down. Ligamental sinus triangular, about as deep as wide, its an-
terior margin vertical to the hinge-line, or slightly descending backward.

Cavity of shells and beaks moderate. Nacre whitish, more or less iridescent;
only in the young specimen with irregular and indistinct radial stria3. Prismatic
border rather narrow, subconcentric with margin, and nowhere noticeably widened,
except very slightly so in the region of the projecting part of the lower margin.
Anterior adductor-scar well impressed, subovate, united above with scar of an-
terior retractor. Anterior protractor-scar united with or separated from that
of the adductor. Posterior adductor-scar faint, subovate, the posterior retractor-
scar forming a triangular upper projection of it. Pallial line subconcentric to the
margin.

According to Wagner, the height would be 57 pr. ct. of the length.

Remarks. — This species has a quite characteristic shape, which varies only
slightly, and the most prominent features are the blunt angle of the lower margin.

604

MEMOIRS OF THE CARNEGIE MUSEUM.

the anterior position of the beaks, the sub-pointed posterior end, and the radial
rib upon the posterior slope. The dark green color of the disk may also be charac-
teristic, but this is liable to fade; in our specimen from Asuncion, a much-worn
shell, the remnants of the epidermis are brown.

IMeasurements.

Localities.

Sex.

Lengtli.

Height.

Diameter.

Beaks.

Santa Rita

cf

36 nun.

22 nini. =61 pr. ct. of L.

13 inni. =36 pr. ct. of L.

at 9.5

mm. =26 pr. ct. of L.

Do

9

62 “

37 “ =59

22 “ =.35

11

“ =18

Limav

?

08 “

41 “ =60

24 “ =35'

16

“ =24

Asuncion

?

70 “

42 “ =60

24 “ =34

16

“ =23

Spix’ figure

48 “

27 “ =56

D’Orbigny’s figure. . . .

=54

= 34

Anatomy. — The soft parts of a young male and a gravid female (collected
June 12) are at hand for study.

The tyiiical Anodonlites-stmGture is observed. The mantle-connection separat-
ing anal and branchial openings is rather long. The Ijranchial opening has ex-
tremely fine papillae. The inner edge of anal and branchial is brown. Palpi of
medium size, semicircular, with a short posterior truncation.

In the gravid female the eggs fill the water-tubes of the inner gill, with ex-
ception of those near the extreme anterior and posterior ends. The water-tubes
are markedly expanded, and the eggs are located in the basal part of the tubes,
and only in the inner coniiiartment (ovisac) toward the inner lamina of the gill.
No larvcU were seen. The outer gills have the usual structure of alternately thicker
and thinner septa, while the septa of the inner gill are more uniform and thicker.
The gills of my female are much torn and injured, so that it was not expedient to
section them.

54. Anodontites hyrioides Ortmann, sp. nov.

Shells: Plate XLII, figs. 3, 4, 5.

Type-locality. — Rio Tapajos, Santarem, Para, Brazil (J. D. Haseman coll.,
December 0-12, 1909). Type-set: Cam. Mus. Cat. No. 61.5829. Six specimens.

Characters of the Shell. — Shell moderately thick, angularly subovate, some-
what olilique, pointed liehind, lower margin convex, forming a blunt angle. Height
62 to 73 pr. ct. of the length. Valves not gaping. Dorsal margin practically
straight, gently descending in front of the beaks. Posterior angle of dorsal margin
obtuse, but quite distinct, wing-like. Anteriorly the dorsal margin forms also a
more or less distinct angle, which is obtuse or almost a right angle. Posterior
margin obliquely descending, straight, or slightly concave in its upper portion,

ortmann: south American naiades.

605

curving sharply around into the lower margin, thus forming the posterior point of
the shell. This point is only a little elevated above the base-line, since the pos-
terior part of the lower margin runs straight forward, being almost parallel to
the dorsal margin (there is some variation in this respect; on the average these
two margins diverge very slightly towards the front). From a point at, or slightly
behind, the middle of the shell the lower margin changes its direction abruptly,
running forward and upward, thus forming a lower blunt angle at about the middle
of the shell. The ascending anterior part of the lower margin is first almost straight,
but then it passes in a curve into the anterior margin, which is narrowly rounded.
The shell appears slightly narrower anteriorly, with the greatest height in the pos-
terior part. The taper of the posterior end is stronger than in A. trigona, and thus
the shell ajipears somewhat oblique.

Valves more convex than in A. trigona, diameter 38 to 46 pr. ct. of the length.
The greatest swelling is towards the beaks, but the beaks are not much elevated
above the hinge-line, so that they appear rather depressed. Location of beaks
at 27 to 31 pr. ct. of the length of the shell. The convexity of the valves is like
that of A. trigona, compressed upon the posterior slope, and flattened in front of
the posterior ridge. Posterior slope with one or two more or less distinct radial
ribs, accompanied by shallow depressions.

Epidermis similar in sculpture to that of .4. trigona, but more frequently
with fine scalariform stripes, chiefly upon the anterior part of the shell, producing
the appearance of fine radial sculpture. Color of eiiidermis from dark or light
greenish and yellowish to light or dark brown; one specimen inclines more toward
olive-green. Most of the specimens are somewhat concentrically banded with
lighter and darker color.

Hinge-line straight behind the beaks, inclining downward in front of them,
but very little so in the youngest specimens. Ligamental sinus triangular, shaped
like that of A. trigona.

Cavity of shell moderate, that of beaks somewhat deeper. Nacre whitish and
iridescent, with indistinct radial striin. Prismatic zone narrow, subconcentric
to the margin. Muscle- and mantle-scars as in A. trigona.

Measurements.

No.

Length.

Height.

Diameter.

Beaks.

Figured.

1

32 nmi.

22 mm

= 68 pr. ct. of L.

14 mm. =44 pr. ct. of L.

at 10 mm. =31 pr. ct. of L.

PI. XLII,

fig. 5.

2

33 “

24 “

= 73

14 “ =42

9 “ =27

.3

41 “

27 “

= 66

16 “ =39

12 “ =29

4

42 “

26 “

= 62

16 “ =38

12 “ =29

PI. XLII,

fig. 4.

5

42 “

29 “

= 69

17 “ =40

12 “ =29

6. . . .

52 “

33 “

= 64

24 “ =46

14 “ =27

PI. XLII,

fig. 3.

606

MEMOIRS OF THE CARNEGIE MUSEUM.

Retnarks. — This species undoubtedly is closely allied to A. trigona, and dif-
fers chiefly in the dimensions. It might be a local variety of it, but since all six
of my specimens are rather uniform in their characters, I take it for a species.

A. hyrioides is an A. trigona, which is higher and shorter, more inflated, has
the outline more sharply angular, and is more oblique. The obliquity is due to

the shortening of the shell and the lower position of the posterior point. The out-

line of our species recalls the shape of the genus Hyria, and hence the name.

55. Anodontites mortoniana (Lea) (1834).

Anodonta mortoniana Lea, Obs. I, 1834, PI. 13, fig. 37.

Anodonta weddellii Hupe, 1857, p. 87, PI. 17, fig. 5.

Anodonta lingulata Hupe, 1857, p. 89, PI. 18, fig. 1.

Anodon weddellii and lingulata Sowerby, XVII, 1868, PI. 20, fig. 80; 1869, PL 23,
fig. 90.

Glabaris ■mortoniana and lingulata Von Ihering, 1893, p. 118, 119.

Glabaris weddelli, lingiilatus, and mortonianus Simpson, 1900, pp. 928, 929.
Anodontites weddelli, lingulatus, and mortonianus Simpson, 1914, pp. 1442-1445.

T ype-locality . — River Parana.

Other Localities. — Santa Ana de Chiquitos, Bolivia (Hupe, weddelli) (situated
about on the divide between the drainages of the Paraguay and the Amazon);
Corumba, Matto Grosso, Brazil (Hupe, lingulata)] Rio Paraguay (Von Ihering);
Lower Parana (Von Ihering).

New Localities. — Mountain creek, Sapucay, Paraguay (S. E. of Asuncion)
(J. D. Haseman coll., April 5, 1909). One male with soft parts. Headwaters of
Rio Paraguay, Santa Rita, Matto Grosso, Brazil (J. D. Haseman coll., June 12,
1909). Two specimens, one a male with soft parts.

Distribution. — This species aiiparently belongs to the Paraguay-drainage and
the Rio Parana below its union with the Paraguay. Von Ihering (1893, p. 114)
does not mention an}^ of the forms belonging to this species from the ujiper Parana.

Remarks as to Aynonymy. — The three forms, mortoniana, weddelli, and lingulata,
are kept as sejiarate species by Simpson, but I do not see any essential differences
between them.

Hupe admits that weddelli is very close to mortoniana, but says that it differs
in three respects: more swollen shell; narrower and more rounded anterior end;
and deeper and larger muscular inqiressions. These differences are not sub-
stantiated by the figures, in fact, we cannot judge as to the first, since no figure
showing the obesity is given. According to the measurements in the text, how-

oetmann: south American naiades.

607

ever, just the opposite is the case, mortoniana being more swollen (40 pr. ct.) than
weddelli (33 pr. ct.). The second difference is not at all correct according to the
published figures; and the third apparently is founded only upon the slight indica-
tion of the muscle-scars in Lea’s figure, and is not essential, anyhow. Besides,
Hupe says that the specific difference is supported by the different distribution
of the two forms. This again is not evident. A. weddelli is from the region of
the divide between the Amazon and Paraguay-drainages, and it may very well be
from the latter. One of our localities is not very far from it. A. mortoniana and
weddelli, indeed, have been united already by Von Ihering (1893, p. 118).

A. lingulata differs from the others only in size and the color of the epidermis.
The latter, on the plate, is dark green, while the text says that it is blackish brown,
and thus we cannot rely on it. The shell of lingulata is more regularly elliptical,
but this is not astonishing when we consider the greater age of this shell. It is
also from the same general region (upper Paraguay) as mortoniana and weddelli.

Very similar forms are found also in the Amazon-drainage: castelnaudi Hupe,
solidula Hupe, amazonensis Lea, and elongala Swainson. But these cannot be
united with mortoniana, since they all are more elongated.

Characters of the Shell. — Shell quite thick and solid. Outline subovate or
nearly subelliptical, bluntly pointed liehind, lower margin gently convex. Height
61 to 63 pr. ct. of the length, falling, in old specimens, as low as 53 pr. ct. Valves
not gaping. Dorsal margin gently convex; the part behind the beaks may be
almost straight; in front of the beaks it descends more or less. Posteriorly the
dorsal margin forms a blunt angle, or may pass almost insensibly into the posterior
margin. There is no distinct angle anteriorly. Posterior margin obliquely descend-
ing, more or less convex, curving into the lower margin and forming with it a dis-
tinct, but rounded, posterior point of the shell, which is situated at a certain eleva-
tion above the base-line, but nearer to the latter than to the line of the upper margin.
Lower margin gentl}^ and rather regularly cuiA'ed, ascending somewhat toward
the posterior end of the shell, and more strongly so in its anterior part, where it
passes in a regular curve into the anterior margin. The anterior part of the shell
is only slightly narrower than the posterior, which is widest (highest) a little behind
the middle of the shell, and then tapers genth" toward the iiosterior point. The
shell is thus transverse, and hardly oblique.

Valves moderately convex, diameter 37 to 41 pr. ct. of the length {weddelli is
more compressed, 33 pr. ct.). Beaks moderately convex, not very prominent above
hinge-line, at 23 to 25 pr. ct. of the length, that is to say, rather anterior. Convexity
of valves rather uniform all over the disk, strongest over the posterior ridge, slightly

608

MEMOIRS OF THE CARNEGIE MUSEUM.

com]ircsscd upon the posterior slope. Posterior slope with one or two blunt
radial ribs, which may be more or less distinct, or almost effaced.

Ejiidermis rather smooth, with irregular concentric lines, which become sub-
lamellar upon the posterior slojie and near the lower margin. Traces of radial
sculpture are iiresent in the shajie of fine scalariform stripes, irregularly disposed
upon the shell, and more or less numerous. Color of epidermis brownish or green-
ish. Upon the disk it may be quite green (dark or light), or it may be darker or
lighter brown, with irregular concentric bands of dark green. The posterior slope
is (in my specimens) always light brown, with or without greenish tints. No
color-rays are seen.

Hinge-line gently curved. In the younger specimens the part behind the beaks
is straight; in older ones it is genth" curved, and curves down more distinctly under
the beaks. Ligamental sinus triangular, about as deep as wide, its anterior margin
vertical to the hinge-line.

Cavity of shell and beaks moderate. Nacre whitish, iridescent, only in the
youngest specimen with faint traces of radial striae. Prismatic zone very narrow
(comparatively widest in the young), subconcentric to the margin. Anterior
adductor-scar well im])ressed, chiefly in the older shells, subovate, united above
with the anterior retractor-scar. Anterior protractor-scar united with, or free
from, adductor-scar (this may be different in the right and left valves of the same
shell). Posterior adductor-scar less impressed, subovate, the posterior retractor-
scar forming an up])er triangular projection of it. Pallial line subconcentric to
lower margin.

Measurements.

1 Leiigtli.

Ileiglit.

Diameter.

Beaks.

Santa Rita. . . 52 mm.

33 mm. = 63 pr. ct. of L.

20 mm. = 3S pr. ct. of L.

at 13 mm. = 25 pr. ct. of L.

Sapucav 54 “

33 “ =61

20 “ =37

13 “ =24

Santa Rita. . . fi!) “

42 “ =61

28.5 “ =41

16 “ =23

Lea’s measurements for jnortoniana give for the height 53 pr. ct. and for the
diameter 40 jir. ct. of the length. Thus the height is less here, but I think this
is due to the greater age of this specimen. My largest siiecimen has a peculiar
shape: the posterior end of the shell is drawn down, so that the lower margin is
almost straight. This, however, undoubtedly is an individual character, since the
growth-lines indicate that the young shell had the normal shape.

The measurements for Hope’s shells are as follows:

Length.

Height.

Diameters.

WcddclU . .
LinquUdn . .

.

66 mm.

100 “

33 mm. = 53 pr. ct. of L.

?

22 mm. = 33 pr. ct. of L.

40 “ =40

ortmann: south American naiades.

609

The height of lingulata is given as 92 mm., but there surely is a mistake in
this statement. According to the figure, it would be 53 pr. ct. of the length.

Remarks. — The characters of this species are its rather regular, subovate,
or subelliptical, outline, with a blunt point behind, somewhat elevated above the
base-line. The shell is not very long and of considerable thickness. The jieculiar
green in the color of the epidermis is remarkable, but is not always present.

Anatomy. — The soft parts of two males are at hand for examination.

The structure is typical of the genus. The jiapillae of the branchial opening
are very small. The inner edge of the anal and branchial is brown. The palpi
are of medium size, semicircular, with a short posterior truncation, forming the
posterior margins, which are not united. The septa of the gills are irregularly
stronger and weaker.

56. Anodontites hasemani Ortmann, sp. nov.

Shells: Plate XLII, figs. 6, 7.

Type-localily. — Headwaters of the Rio Paraguay, Santa Rita, Matto Grosso,
Brazil (J. D. Haseman coll., June 12, 1909). Type-set: Cam. Mus. Cat. No.
61.5832. Four specimens, among them two males and a gravid female with soft
parts.

Characters of the Shell. — Shell moderately thick, outline subovate, narrower
in front, broader (higher) behind, not pointed. Height 63 to 67 pr. ct. of the length.
Valves not gaping. Dorsal margin gently curved, the ]iart behind the beaks almost
straight; anteriorly to the beaks it descends distinctly, and passes into the anterior
margin in an indistinct, obtuse angle. At the posterior end the upper margin forms
a more distinct, obtuse angle. Posterior margin oliliquely descending, gently
convex, broadly curving around at the posterior end into the lower margin. The
latter is gently curved, and runs forward and upward, being almost straight in
the anterior part, finallj^ curving u}i into the anterior margin. Thus the shell is
distinctly narrowed anteriorly, the greatest height lieing located at about the be-
ginning of the posterior third of the shell, and the shell being somewhat oblique.

Valves rather convex, diameter 42 to 47 ]n*. ct. of the length. Beaks rather
inflated, but only moderately elevated above the hinge-line, located at 27 to 33
pr. ct. of the length. Convexity of valves rather regular, greatest over the posterior
ridge, which is indistinct, very slightly compressed upon the posterior slope, with-
out perceptible flattening upon the sides of the disk. Posterior slope with indistinct
and faint traces of one or two radial ribs.

Epidermis very slightly shining, with crowded, irregular, fine, subconcentric

610

MEMOIRS OF THE CARNEGIE MUSEUM.

lines, becoming lamellar upon the posterior slope and near the lower margin. There
are hardly any traces of radial sculpture. Color olive-brown, rather uniform.
The smallest specimen has a trace of a dark radial ray u])on the posterior slope,
otherwise there are no rays, nor indications of growth-rests.

Hinge-line ipion the whole gently curved, but the part behind the beaks is
practically straight, chiefl}^ in the younger specimens. The part in front is decurved,
but in one specimen it distinctly curves u]) again at the anterior end. Ligamental
sinus triangular, hardly as dee]i as wide, its anterior margin in the largest specimen
being vertical, in the others slightly descending backward.

Cavity of shell and beaks rather deep, corresponding to the obesity of the
valves. Nacre blueish white, with purple and green iridescence, and with very
faint radial striae in the younger specimens. Prismatic zone quite narrow, sub-
concentric with the margin. Anterior adductor-scar not very deeply impressed,
subovate, united with the scar of the anterior retractor; scar of anterior protractor
separated from it more or less completely. Posterior adductor-scar faint, sub-
ovate, with an upper triangular jirocess formed by the posterior retractor. Pallial
line subconcentric with the margin.

Measurements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Figured.

1

d'

45 mm.

.30 mm. =07 pr. ot. of L.

19 mm. =42 pr. ct. of L.

at 15 mm. =33 pr. ct. of L.

PI. XLII, fig. 0.

2

d

57 "

30 “ =03

24 “ =42

17 “ =32

3

9

02 “

39 “ =03

28 “ =45

17 “ =27

PI. XLII, fig. 7.

4

?

00 “

43 “ =05

31 “ =47

18 “ =27

Remarks. — I cannot find among the described sjiecies any one which agrees
with the ]iresent form. Its chief feature is the broadly rounded posterior end.
In this character and in the oblicpiity it resembles the species of the patagonica-
group, but it differs from them in the iirismatic border, which is narrow. I have
placed this species in the trigoyia-group, although it has not the subpointed pos-
terior end, because I could not conveniently place it anywhere else. Of other
species, only A. ohtusa resemlffes it to a degree, but the latter is somewhat shorter
and higher (Height 64 to 69 pr. ct. of L.), and not distinctly oblique, with the
upper and lower margins more nearly })arallel. Besides, A. ohtusa has a peculiar
color-pattern.

Anatomy. — Soft parts of two males and one gravid female at hand.

The structure is of the normal Anodontites-type. Papillie of the branchial
opening very small. Gills with the septa well developed. In the gravid female,
the inner gills have the usual marsupial structure, with a swelling at the insertion
of the sei)ta on the outer (])rimary) limb of the gill. The water-tubes contain

ortmann: south American naiades.

611

eggs in their inner compartments, except at the anterior and posterior ends of
the gill; the other compartments forming secondary water-tubes. The septa
of the ovisacs are somewhat stretched out, and the gill is slightly distended. I
was unable to find fully developed larvae (lasidia?). All eggs are small, round,
globular masses of cells (morula-stage), enclosed in a rather tough membrane.
In the males no regular alternation of stronger and thinner septa could be noticed,
but all septa are rather uniform. The male character, however, has been positively
established by microscopic examination (absence of swellings of the septa).

5. Group of Anodontites patagonica.

Simpson, 1914, p. 1403.

Shell strongly oblique, subovate or subrotund, not pointed behind. Epidermis
more or less shining, but here and there with wrinkles, or covered all over with
concentric striae. Prismatic liorder rather wide and unequal, being much wider
along the anterior lower margin than at the anterior and jiosterior ends.

The prismatic border forms the most prominent feature of this group. I
possess a good number of specimens belonging to the grou]), and this character is
always present, so that we must regard it as of taxonomic value. In addition
the shell is here distinctly oblique with a rounded end.

Key to the Forms at Hand.

a I. Shell moderately inflated, diameter generally between 40 and 50 pr. ct. of the length, rarely less.

Outline strongly oblicpie, moderately elongate, sometimes subrotund, height from about 65 to
over 80 pr. ct. of length.

bi. Shell thick, rather more elongated. Nacre whitish A. patagonica.

62. Shell thinner, snbrotund or elongated. Nacre often red /I. patagonica rubicunda.

0 2. Shell compressed, diameter 36 pr. ct. or less. Outline obliquely ovate, rather elongated, height not
over 66 pr. ct.

bi. Prismatic border wide and unequal. Epidermis shining, .strife not sublamellar. . . .A. puelchana.
62- Prismatic border narrower, but unequal. Epidermis not very shining, with the strife often
sublamellar A. iheringi.

57. Anodontites patagonica (Lamarck) (1819).

Anodonta patagonica Lamarck, 1819, j). 88; Encyclop. Method., II, 1827, p. 147,
PI. 203, fig. 1.

Anodontites patagonicus Simpson, 1914, p. 1403.

Reinarks as to Synonymy. — The synonymy of this species has lieen given by
Simpson, but it needs certain additions and corrections. The following references
should be added first of all :

Anodonta latemarginata and uruguayensis Corsi, 1901, p. 454, 458.

612

MEMOIRS OF THE CARNEGIE MUSEUM.

Anodontites patagonicus Haas, 1916, pp. 25, 54.

As we have seen above, Anodon trapezeum Spix (1827) should be stricken from
the list of synonyms. This is a different species, differing chiefly by the narrow
lirismatic border. The other references given by Simpson all certainly belong here,
and the following names have been used for this species :

Anodonta latomarginatn Lea (1834).

Anodoyita memhranacea D’Orbigny (1843).

Anodonta solida Kuester (1853).

Anodonta Uruguay ensis Lea (1860).

Anodonta sinuosa Clessin (1873).

Anodonta. serpentina Ci.essin (1876).

In addition:

Glaharis bergi Von Ihering (1893 p. 118), introduced for sinuosa Clessin.

But the following forms also lielong here as synonj^ms:

Anodon crassus Swainson (1823); Simpson, 1914, p. 1406. Simpson says: this
s]iecies is “close to A. patagonicus, but I have never seen a specimen of that
species quite so elongated or so })entagonal as the figure.” The dimensions
are: L. 80 mm., H. 52 mm. = 65 pr. ct. of length. Diameter 32 mm. = 40 pr. ct.
of length. These figures agree well with the dimensions of my series of speci-
mens from San Isidro, some of which are even more elongated (height falling
as low as 60 pr. ct.

Anodonta wymarii Lea (1860); Simpson, 1914, j). 1407, who says: “More elongated
and more richly colored than A. patagonicus." The elongation is even greater
than in A. crassus (Height = 63 pr. ct. of length), but it still remains within
the limits of variation of my set from San Isidro. Some of the latter un-
doubtedly are wyniani in every respect. The color is no reliable character,
being very variable in A. patagonica.

I am unable to form an opinion as to A. sirionos D’Orbigny (1835) (= ferrarisi
D’Orbigny, 1835). It certainly is nearly related to A. pcdagonica, but, according
to the original descrijition, is distinguished by a rough and concentrically and
lamellarly striate epidermis. I have no specimens corresponding to this form.
Type-locality. — Rio de la Plata and Patagonia.

Other Localities. — Rivers of Uruguay between Montevideo and Buenos Aires
(D’Orbigny, m.emhranacea) •, Rio Miguelete, Montevideo, Uruguay (Haas); Arroyo
S. Jose, Uruguay (N. W. of Alontevideo) (Corsi, Uruguay ensis)] Uruguay River
(Lea, uruguayensis, wymani)] Rio de la Plata (Haas); Rio de la Plata, Buenos
Aires, Argentina (D’Orbignj^ m.emhranacea)] Rio Parana (Lea, latomarginata) ]

ortmann: south American naiades.

613

Rio Parana up to sixty miles above Corrientes, Argentina (D’Orbigny, mem-
hranacea) .

Localities Represented in the Carnegie Museum. — Rio Parana (Hartman collec-
tion). One specimen. In pond along banks of Rio Negro, Santa Isabel, Urugua}''
(J. D. Haseman coll., February 11, 1909). One specimen. Arroyo Miguelete,
Montevideo, Uruguay (J. D. Haseman coll., February 17, 1909). One specimen.
Rio de la Plata, San Isidro, 20 km. N. of Buenos Aires, Argentina (A. Windhausen
coll., January, .1917). Twenty specimens with soft parts, males and females.

Distribution. — La Plata system, from the mouth near Buenos Aires u]) the
Parana to the province of Corrientes in Argentina; also in the Rio Uruguay and the
Rio Negro, and the tributaries of the La Plata in the Banda Oriental of Uruguay.

Characters of the Shell. — Shell rather large (length up to 100 mm. and over),
rather thick and solid. Outline very obliciue, subovate, longer or shorter, height
60 to 74 pr. ct. of the length. Young shells probably higher and more subrotund,
according to the growth lines in older shells, but such shells have not been observed
(the smallest shell is 75 mm. long). Valves not gaping. Dorsal margin straight
or gently curved, generally with a well marked, blunt posterior angle, while the
anterior angle is less distinct, often quite rounded. Posterior margin obliquely
descending, generall}^ almost straight, rarely gently convex, and very rarely a
little concave, curving broadly and regularly into the posterior lower margin,
generally forming no angle or posterior point, but sometimes with a trace of it-
The anterior part of the ventral margin is strongly ascending and always more or
less gently curved, and often almost straight, curving u]) into the anterior margin,
which is narrowly rounded. Thus the anterior end of the shell appears much nar-
rower than the broadly rounded posterior part, rendering the shell decidedly oblique.

Valves moderatel}'^ swollen, diameter 38 to 44 iir. ct. of length, rarely slightly
less. Beaks slightly convex, and only little elevated above the hinge-line, located
at 28 to 34 pr. ct. of the length. Disk moderately convex, more strongly so over
the middle part and the posterior ridge; iiosteriorly compressed. Posterior ridge
very indistinct, but often there is a radial rib uiion the posterior slope, sometimes
accompanied by a furrow. The anterior end of the shell is also slightly compressed.

Epidermis rather smooth, but with tine, irregular, concentric striie, almost
effaced in the middle of the disk and towai’d the beaks, more distinct toward the
margins, where they may become sublamellar. Very fine radial, oblique, or
reticulated wrinkles may be present between the striae. Scalariform radial stripes
are generally absent, but in some specimens slight traces of them are seen. Color
of epidermis dark greenish-olive to various shades of brown. Generally there is

614

MEMOIRS OF THE CARNEGIE MUSEUM.

more green toward the beaks, more brown toward the margins, but the green
may extend over nearly the whole shell, or may be absent. Upon the posterior
slope there may be a few indistinct, dark green rays, and the green color near
the beaks often has a mottled character, consisting of irregular, darker and lighter
concentric bands, and, rarely, of indistinct radial rays or blotches. In some
specimens, the anterior and jiosterior parts of the shell are different in shade, the
posterior being darker (more greenish), the two shades being separated rather
sharjily by a radial line. Growth-rests few, and rarely distinct.

Hinge-line nearly straight, or very gently curved behind the beaks. In front
of them it descends more or less distinctly, and may ascend again at the anterior
extremity, thus liecoming slightly sinuate. Ligamental sinus triangular, some-
what varialile, Imt generally deeper than wide, chiefly in older shells, its anterior
margin mostly vertical to the hinge-line, and its lower ])oint curved more or less
forward. Often the anterior margin has an S-sha]ied curve.

Cavity of shell and beaks moderate. Nacre whitish, iridescent, but none of
my specimens has red tints. Radial striations are present, but indistinct in older
shells. Prismatic border broad, unequal in width, broadest along the ascending
anterior lower margin, narrower in front, and narrowing rather suddenly at the
beginning of the posterior i)art of the lower margin. Its color is grayish or yellowish
green, or grayish or yellowish white. Anterior adductor-scar well impressed,
subelli])tical, united above with that of the retractor, or sometimes incompletely
separated. Anterior jirotractor-scar generally well sejiarated from the adductor-
scar. Posteilor adductor-scar less imjiressed, sul^ovate, with an upper triangular
])rojection formed by the posterior retractor-scar. Pallial line distinct, subcon-
centric with the shell-margin, and thus more closely aiiproaching the prismatic
liorder in the anterior part of the lower margin than before and behind it. No
dorsal scars.

Measurements.

Locality.

No.

Sex.

Length.

Height.

Diameter.

Be.aks.

San Isidro

10

d'

75 nun.

53 nun

= 71 pr. c

t. of L.

30

nun. =40 pr. c

t. of L.

at 25 nun

= 33 pr. ct. of L.

Do

14

d

87

50

= 64

“

38

“ =44

“

29 “

= 33

Do

lU

9

89

62.5 “

= 70

“

39

“ =44

“

25 “

= 28

Do

2

9

91

67

= 74

“

36.;

" =40

“

27 “

=30

Do

20

9

93.5 “

56

= 60

“

39

“ =42

“

26 “

=28

Do

8

(T

103

66.5 “

= 65

37

“ =36

‘ ‘

31 “

= 30

Montevideo

?

77

51

= 00

“

30

“ =39

26 “

= 34

Santa Isabel

98

07

= 68

“

42

“ =43

“

33 “

= 34

Previous Measurements.

D’Orbigny (memhranacea) ....

90

=73

= 33

Simpson (pHtayonica)

81

59

“ =73

32

“ =40

Do.

87

57

“ =00

37

“ =43

Simpson (crassa)

80

52

“ =65

32

“ =40

Simpson (mjmuni)

80

50

“ =63

30

“ =38

Lea (figure of u»/mani)

87

55

“ =03

33

“ =38

ortmann: south American naiades.

615

All of these iirevious measurements fall within the range of variation of my
series, with the exception of the diameter given by D’Orbigny, which is somewhat
lower (33 pr. ct.) than any of my figures (lowest 36 pr. ct.).

Re7narks. — This species is very variable in shape, being sometimes higher,
sometimes more elongated. There is no indication that the shape is connected
with sex. Since no very young specimens are at hand, and never have been ob-
served, we do not jiositively know anything about their shape, but from the growth-
lines of the old specimens it is seen that the young shell generally must have been
comparatively higher; the shell grows, with advancing age, more in the direction
of the posterior end, so that the longitudinal diameter increases more than the
vertical.

There is no doubt in my mind that A. crassa and wijmani belong here, and that
they represent specimens which are a little more elongated than the average; but
among the set from San Isidro, I have specimens which represent even greater ex-
tremes than these.

A. patagonica is rather thick and solid, but varies also in this respect. The
specimen from Montevideo at hand is not quite as thick as the others, but agrees
with them in other respects, being also rather elongated. Just such specimens
induce me to regard the next form {rubicwnda) as a variety of patagonica.

Anatomy. — Soft parts of eight males, one barren, and eleven gravid females
have been investigated. The breeding season is January.

The anatomy of latomarginata and wymani has been previously described by
Lea (Obs. X, 1863, pp. 391, 394) as far as the superficial characters of the gills,
the palpi, and the branchial and anal openings are concerned.

Von Ihering (1891, p. 480) describes the eggs and lasidia of Glaharis wymani,
the former being 0.09 mm. in diameter, the latter 0.086 mm. long. However, since
he says that his specimens are from Rio Camaquam, I am not sure that they have
been properly identified. They might belong to A. iheringi (Clessin).

Anal opening entirely open, separated from the branchial b}^ a connection of
the mantle-margins. Inner edge of anal smooth above, very finely crenulated
near the lower end. Branchial opening with small papillae. Pal])! nearly semi-
circular, longer than high, behind abnqitly truncated, the posterior margins con-
nected at base.

Gills rather wide, the inner much wider than the outer in front, its anterior
end immediately behind the palpi, and attached along the whole space between
the palpi and the anterior end of the outer gill. Inner lamina of inner gill entirely
connected with abdominal sac. Septa of the gills well developed. In the non-

616

MEMOIRS OF THE CARNEGIE MUSEUM.

marsu])ial gills tliej" are irregularly alternating. In the female, the inner gill is
marsupial nearly throug^ioiit its whole length. The septa are stronger and equal,
having near the outer lamina a swelling indicating ridges projecting into the lumen
of the water-tubes. When charged the egg-masses occupy only the inner com-
imrtments, forming ovisacs, which are somewhat distended, while the outer com-
partments do not change, and apparently serve as (secondary) water-tubes. Eggs
very small. 1 did not see any larvse. Even in a specimen which had the gills
only jiartially charged (anteriorly), and which might have been discharging, the
eggs consisted only of a globular mass of cells enclosed in a membrance.

57a. Anodontites patagonica rubicunda (Lea) (1860).

Diagram of soft parts: Text-fig. 3, p. 458.

Section of gills: Plate XLVllI, fig. 8.

Anodonta rubicunda Lea, Obs. X, 1863, PI. 46, fig. 299; Corsi, 1901, p. 455.
Anodotda pazii Lea (1866) Obs. XII, 1869, PI. 36, fig. 87.

Anodon rubicundus Sgwerby, XVII, 1870, PI. 30, fig. 118.

Glabaris rubicunda Pilsbry & Rush, 1896, p. 81; Simpson, 1900, p. 913.

Glabaris latomargincdus felix Pilsbry, 1896, p. 563, PI. 26, fig. 8.

Glabaris pazii Simpson, 1900, p. 918.

Anodontites latomarginaia felix Simpson, 1914, p. 1405.

Anodontites pazi Simpson, 1914, p. 1408; Haas, 1916, pp. 30, 55.

Anodontites rubicundus Simpson, 1914, p. 1409.

Type-locality. — Uruguay River.

Other Locatities. — Rio de la Plata, Colonia, Uruguay (Pilsbry & Rush, rubi-
cunda) ( Pilsbry, felix) ; Uruguay River, Paysandu, Uruguay (Pilsbry & Rush) ;
Uruguay River, Salto Oriental, Uruguay (Haas).

ATic Locatities. — Rio Uruguay (in mud), Uruguayana, Rio Grande do Sul,
Brazil (J. D. llaseman, February 5, 1909). Twelve specimens, eleven of them
with soft parts, males and females. In pond along banks of Rio Negro, Santa
Isabel, Uruguay (J. D. llaseman coll., February 11, 1909). Three specimens.
Rio Ibicuhy, Gacequy, Rio Grande do Sul, Brazil (J. D. Haseman coll., February
1, 1909). One male with soft parts. Rio Gacequy (in sand), Gacequy, Rio Grande
do Sul, Brazil (J. D. Haseman coll., February 2, 1909). One female with soft
parts.

Distribution. — Known only from the Uruguay and the Rio de la Plata below
tlie mouth of tlie Uruguay, and from the Rio Negro in Uruguay and Rios Ibicuhy
and Gacequy in Rio Grande do Sul, these being tributaries of the Uruguay.

ortmann: south American naiades.

617

Closely allied to A. patagonica, but differing in being somewhat smaller (maxi-
mum length 84 mm.), with a thinner shell. The outline of the shell is more nearly
and more frequently subrotund, and the diameter ranges higher (up to 50 pr. ct.).
The nacre very often, but not always, has reddish or purple tints.

Measurements (Specimens from Uruguayana).

No.

Sex.

Length.

Height.

Diameter.

Beaks.

1

c?

47.5

mm.

36.5

mm. =77 pr. ct. of L.

19.5

mm. =41 pr.

ct. of L.

at 17

mm. =36pr.ct.otL.

b

9

53

44.5

“ =84

24.5

“ =46

19

“ =36

r2

9

59

42

“ =71

22

“ =37

20

“ =34

d

9

62

50

“ =81

31

“ =50

24

“ =39

3 . . . . .

&'

66

49

“ -74

27

“ —41

99

“ -33 “

e

9

75

58

“ =77

33

“ =44

22

5 “ =30

4

9

84

59

“ =70

35

“ =42

29

“ =35

Older Measurements.

Simpson,

rubicunda.

59

51

“ =86

30

“ =51

(Type)

Do.

77

61

“ =79

36

“ =48

Simpson,

pazi

60

43

“ =72

26

“ =43

“ (Type)

Do.

53

38

“ =72

26

“ =49

Pilsbrv, fdix

49

35

“ =71

IS

“ =.37

Do.

53

38

“ =72

20.5

“ =39

Remarks. — All the previous measurements fall within the range of variation of
my specimens, only the type of Lea’s rubicunda is higher, shorter, and more swollen
than any of my specimens. Thus we are to regard the type of rubicunda as an
exceptional specimen, which is also peculiar in the fact that the posterior margin
forms nearly a right angle with the upper margin. All these peculiar features are
correlated, however. The normal type of this form is better represented by Lea’s
pazi] yet there are specimen in my material which distinctly approach the rubicunda-
type.

It is unknown whether this form ever reaches the size of A. patagonica, and
in the absence of young specimens of the latter it is impossible to compare them
directly. But it seems that the more nearly rotund shape is rather a juvenile
character, although there are young specimens in my material which are more
elongated. From the measurements given it is seen that the height ranges from
70 to 84 pr. ct. in my material, reaching 86 pr. ct. in Lea’s type of rubicunda, while
it ranges from 60 to 74 pr. ct. in patagonica. The diameter in rubicunda is from 37
to 50 pr. ct. (51 pr. ct. in the type), while in patagonica it is from 36 to 44 pr. ct.

Clearly height and diameter are correlated, a shorter and higher shell being
also more swollen than a more elongate one. The slight difference in the location
of the beaks (30 to 39 pr. ct. in rubicunda, 28 to 34 jir. ct. in patagonica) is also
connected with this. Finally it may be that all these differences in the shape are
connected with age, and this may be true also of the thickness of the shell. How-

618

MEMOIRS OF THE CARNEGIE MUSEUM.

ever, niy largest specimens of rubicunda differ in this from the smallest specimens
of patagunica (of about the same size), but not very strikingly.

Much stress has been laid in the original descriptions of A. rubicunda and
pazi upon the red color of the nacre. According to my material this is extremely
variable. In some specimens the color does not differ from that of A. patagonica,
being whitish. In others it is more or less tinted with red or purple, and then also,
generally, the prismatic border is didl purplish gray. But this also is not a constant
character.

Pilslny’s felix is said to be characterized by its light, yellow-green epidermis,
which has radiating or irregularly zig-zag lines of green. I see traces of this in
some of my specimens from Uruguay ana and Santa Isabel, and there are also here
and there black markings on the inside along the pallial line and near the muscle-
scars (and elsewhere), mentioned by Pilsbry as occurring in felix. These are in-
dividual characters.

The epidermis of rubicunda is much as in patagonica, i.e., somewhat lamellar
near the margins, smoother in the middle. Scalariform stripes are sometimes
indicated, but are mostly absent. The color in younger specimens is generally dark
green all over the shell, but a few are brownish. Those with intense red tints
on the inside are sometimes a little reddish brown on the outside. The largest
specimen in the color of the epidermis is much like patagonica, brownish, inclining
to olive toward the beaks.

D’Orbigny says of patagonica {niembranacea) that it is very variable; that it
is larger and thicker in the larger rivers; more elongated in lakes; and that it be-
comes reddish on the inside in small rivers. My specimens support the latter
observation, for they all are, so far as they may be called rubicunda and have red
nacre, from comparatively small rivers (Rio Negro, upper Uruguay, Ibicuhy, and
Cacequy). But, of course, this requires further study and more material.

Anatomy: The soft parts of six males and seven females are at hand, one of the
latter gravid (February 5).

The anatomy of two males of pazi has been described by Lea, but only the
major features have been mentioned. According to my material the soft parts
agree entirely with those of the typical patagonica. In the gravid female the
swelling of the charged marsupium is not considerable. The small eggs hang
loosely together, and easily fall ajiart. No fully developed larva3 could be found.
A diagrammatic figure of the soft parts is given in text-figure 3 (p. 458), and the
structure of the gills of a barren female in cross-section is shown on PI. XLVIII,
fig. 8.

ortmann: south American naiades.

619

58. Anodontites puelchana (D’Orbigny) (1835).

Anodonta puelchana D’Orbigny, 1843, p. 620, PI. 79, figs. 7-9.

Glabaris puelchanus Simpson, 1900, p. 921.

Anodontites puelchanus Pilsbry, 1911, p. 609.

Simpson (1914, p. 1420) unites this with A. limnoica D’Orbigny, but the latter
has a very narrow prismatic border. According to my observations, the width
of this border is a very constant and important character. Simpson places this
species in the group of crispata; but there is not the slightest trace of the character-
istic sculpture of the epidermis of this group. That Simpson has entirely mis-
understood this species is shown by the fact that he unites A. obtusula Hupe with
it. The latter is a species which is not at all oblique.

Type-locality . — Marsh of San Xavier on Rio Negro, above Carmen de Pata-
gones, Argentina.

Additional Locality. — Twelve leagues from Chichinal, Patagonia (on Rio Negro)
(Pilsbry).

New Locality. — Rio Limay, Patagonia (drainage of Rio Negro) (W. Israel
donor). Two specimens with soft parts, male and female.

Distribution. — Known only from the Rio Negro-drainage in Patagonia. Von
Ihering (1893, p. 118) lists it as from the lower Parana, but without substantiating
this record.

According to D’Orbigny ’s description and figure, this species is an obliquely-
ovate, much compressed, thin shell, with shining epidermis, and wide prismatic
border. It is surely related to patagonica, being, however, not so high and more
elongated (height 57 to 65 pr. ct. of length, while patagonica has the height 60 to
84 pr. ct. The diameter is about 31 pr. ct. while in patagonica it never falls below
33 pr. ct. and only rarely below 40 pr. ct.

The two specimens before me have the wide prismatic border, somewhat
unequal in width; the nacre is whitish with pinkish shades (D’Orbigny says “blanc-
rose”). The epidermis is highly polished and shining; there are some concentric
grooves, but the fine concentric striae are missing upon the greater part of the disk,
and are only slightly developed near the margins. There are, chiefly in my larger
specimen, a few scalariform stripes upon the anterior part of the shell, consisting
of radial bands of short, concentric wrinkles. Upon the posterior slope there is
an indication of a radial rib. Color of epidermis greenish brown, more greenish
toward the beaks, more brownish toward the margins, with very indistinct traces
of green rays upon the posterior slope.

620

MEMOIRS OF THE CARNEGIE MUSEUM.

IMeasukements.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

1

cf

40 mm.

2G mm. = G5 pr. ct. of L.

12.5 mm. = 31 pr. ct. of L.

at 13.5 mm. = 34 pr. ct. of L.

2 . . . .

cf

65

40 “ =G2

20 , “ =31

21 “ =32

D’Orbigiiy .

GO “

= 57

= 31

Anatomy. — The soft parts of the male and female at hand are not in good con-
dition, the female being the better of the two. The usual structure of the genus,
however, could be made out, and no special features require mention.

59. Anodontites iheringi (Clessin) (1882).

Shells: Plate XLIl, fig. 8; Plate XLIII, fig. 1; Plate XLIV, fig. 1.
Anodonta iheringi Clessin, Malakazool. Blaett., V, 1882, p. 191, PI. 4, fig. 5.
Glabaris iheringi Simpson, 1900, p. 919.

Anodontites sirionos ihermgi Simpson, 1914, p. 1408.

Type-locality . — Taguara del Mundo Novo, Rio Grande do Sul, Brazil (system
of Rio Guahyba, N. E. of Porto Alegre).

New Localities. — Rio Jacuhy, Cachoeira, Rio Grande do Sul, Brazil (J. D.
Haseman coll., January 26, 1909). One young specimen, with soft parts, sex not
determined. Rio Vaccahy-Mirim, Santa Maria, Rio Grande do Sul, Brazil (J.
D. Haseman coll., January 29, 1909). Three specimens, with soft parts, two males
and a female.

Distribution. — Guahylia-drainage in Rio Grande do Sul.

This species very closely resembles A. patagonica, but is more elongated and
oblique, so that in outline it is still more like A. puelchana. To the latter species
it is related also in the more coinjiressed shell.

The shell is rather thin, of medium size. The outline is obliquely ovate,
narrower anteriorly, broader (higher) posteriorly. The lower margin ascends in
its anterior portion, and may be gently curved or nearly straight. Valves gently
convex, rather comiiressed. The prismatic border is not so wide as in patagonica
and puelchana, but it is of the same general character, being wider along the an-
terior lower margin. It is of a grayish green color. The nacre is silvery white,
highly iridescent, often greenish or grayish discolored. Ligamental sinus about
as deeji as wide, its anterior margin vertical, with the point bending forward.

The chief difference from patagonica, and especially from puelchana, is in
the texture and color of the epidermis. The latter is brownish or brownish olive,
not very shining, covered with close, subconcentric stria3, sublamellar near the
margins. Scalariform stripes are distinct upon the anterior half of the shell.

ortmann: south American naiades.

621

consisting of radial bands of short, subconcentric wrinkles. These stripes may
also be seen upon the posterior section of the shell, but less frequently and less
distinctly.

Measurements.

Location.

6

y.

a>

02

Length.

■

Height.

Diameter.

Beaks.

Figured.

Cachoeira . . .

'?

22.5 mni.

14.5nmi. =64 pr.ct. of L.

8 mm. =35 pr. ct. of L.

at 6.5 mm. =29 pr. ct. of L.

Santa Maria.

1

d'

53 “ 35 “ =()()

18 “ =.34

17 “ =32

PI. XLII, fig. 8.

Do.

2

d

56

36.5 “ =65

20 " =36

18 “ =32

Pi. XLIII, fig. 1

Do.

3

9

69 “ ,43 “ =62

25 .“ =36

21.5 “ =31

PI. XLIV, fig. 1

Simpson’s measurements are: Length 61 mm.; Height 39 mm. = 64 pr. ct. of L.;
Diameter 22 mm. = 36 pr. ct. of Length.

Remarks. — The original description and that given by Simpson are scarcely
sufficient to enable the species to be recognized, but none of the characters given
conflict with those exhibited by my specimens, and the measurements apply very
well. Since the original A. ihcringi comes also from the same river-system (Guahy-
ba) as my specimens, I have no doubt that we are dealing with this species.

Anatomy. — Two males, and one female are at hand for study.

The soft parts are absolutely identical with those of A. patagonica.

6. Group of Anodoniites trapesialis.

Shell strongly oblique, subovate to subtrapezoidal, narrowly rounded, or
bluntly pointed behind. Epidermis shining, only here and there, and not always,
with scalariform radial bands of wriidvles. Prismatic border narrow and of equal
width.

A rather well defined group, distinguished also by the large and comparatively
thin shell: the largest species of the genus belong here. Usually we discover that
the valves are gaping anteriorly, and sometimes also posteriorly. Whether this
is connected with the anatomical structure, or with habits, is not known. There
is hardly a question that the genus Leila is descended from forms belonging to this
group.

Von Ihering has given a partial key to the species (1890, p. 157), which I
have used to great advantage. But great difficulties arise in distinguishing the
species, and young individuals are often very hard to correlate with older ones.
Although I possess good material representing this group, it includes only com-
paratively few forms, so that I am unable to form a definite opinion as to those which
are not represented. I treat my specimens here under the names to which they
seem to belong, without going into detail as to their affinities and relationships.

622

MEMOIRS OF THE CARNEGIE MUSEUM.

60a. Anodontites TRAPESiALis ANSERiNA (Spix) (1827).

Simpson, 1914, p. 1430.

New Locality. — Rio Paraguay, Corumba, Matto Grosso, Brazil (H. H. Smith
coll.). Three right valves.

I do not jiropose to go into a lengthy discussion of this form. I merely desire
to say that two of my specimens agree very well in shape and size, with the figures
given by Spix, while the third more resembles Sowerby’s figure (PI. 31, fig. 125).
They vary slightly among themselves in the degree of the taper of the anterior
end. In all of them the hinge-line is very gently curved, but not sinuous (as in
A. exotica).

The original A. anserina is from the Rio Solimoes, in the province of Amazonas.

606. Anodontites trapesialis scripta (Sowerby) (1867).

Simpson, 1914, p. 1430.

Simiison unites with this A. bahiensis Kuester (Von Ihering, 1890, pp. 153,
157; 1893, p. 115; 1910, p. 138).

No exact localities are known. A. bahiensis, which is given from Bahia, is
not from that region, but, according to Von Ihering (1910) from the upper Amazon
and Ecuador. Von Ihering (1890, p. 152) thinks, on the other hand, that scripta
is identical with exotica.

New Locality. — Rio Ribeira, Iguape, Sao Paulo, Brazil (Haseman collection,
collected by Richardo Krone). Two odd valves, right and left.

My specimens agree well with Sowerby’s (PI. 4, fig. 9) in shape and size. One
of them has the characteristic zig-zag black markings on the inside, while in the
other one these are found only near the posterior adductor-scar. However, the
color of the epidermis in both is not brownish, but green, darker upon the posterior
slope. Vei\y obscure green rays are seen on some parts of the disk. Since Simpson
says that the color of the epidermis varies to the ‘‘ordinary” (green) color, I do
not think that this prevents the union of our specimens with scripta. My left
valve has anteriorly two short scalariform stripes, barely visible; in the right valve
such are entirely absent.

JMeasuhements.

Lengtli.

lleiglit.

Diameter.

Beak.s.

Left

Riglit ....

154 mill.
160 “

03 mill. = 60 jir. ct. of L.
96 “ =60

50 mm. = 32 pr. ct. of L.

1 48 “ =30

at 61 mm. = 40 jir. ct. of L.
60 “ =38

ortmann: south American naiades.

623

61. Anodontites moricandi (Lea) (1860).

Simpson, 1914, p. 1439.

Synonym :

Anodonta hertwigi Von Ihering, 1890, p. 150, PI. 9, fig. 7; 1910, p. 138.

Von Ihering (1910, p. 138) regards Anodon radiatus Spix (1827) as the young
of this species, and in that case, of course, the specific name radiata should supersede
that of moricandi. However, I am not convinced that this is right, for those of
our specimens which are nearly of the size of Spix’ original, do not agree with it
in shape. They do not have the strongly convex lower margin, and the relative
dimensions, chiefly the height, also do not fit. According to Von Ihering, the
measurements of radiata are: Length 70 mm.; Height 36 mm. = 51 pr. ct. of L.;
Diameter 20 mm. = 29 pr. ct. L.

Type-locality . — Bahia, Brazil.

Other Localities. — Rio Sao Francisco, Villa Nova, Sergipe, Brazil (Von Iher-
ing); Rio Paraguassu, Bahia, Brazil (Von Ihering); Rio Pardo, Bahia, Brazil (Von
Ihering).

New Locality. — In a lagoon of Rio Parahyba, Campos, Rio de Janeiro, Brazil
(J. D. Haseman coll., June 14, 1908). Six specimens.

Distribution. — Streams of eastern Brazil, from mouth of Rio Sao Francisco in
Sergipe southward to Rio Parahyba in Rio de Janeiro. Known from the rivers
Sao Francisco, Paraguassu, Pardo, and Parahyba.

My specimens agree well with this species. What is regarded as one of its
essential characters, the flattening of the disk on the sides, is developed only in
larger specimens. My largest specimen shows it distinctly, although not as strik-
ingly as Lea’s figure. In the smaller ones this is less evident, but even in these
the greatest diameter of the shell is in about the middle of the length, not much
behind the beaks.

Measurements.

Length.

Height.

Diameter.

Beaks.

30.5 mm.

17

mm. = 56 pr. ct. of L.

10

nini

. = 32 pr. ct. of L.

at 1 1

mm. =36 pr. ct. of L.

49

28

“ =57 “

17.5

“

= 36

18

“ =37

75

45

“ =60

25

a

= 33

24

“ =32

78

44.5

“ =57

23.5

a

= 30

25.5

“ =33

89

50.5

“ =57

25.5

(

= 29

29

“ =33

98

57

“ =58

30

= 31

34

“ =35

104 “

61

“ =59

32

((

= 31

30

“ =29

(Lea’s fig.)

117 “

70

“ =60

35

u

= 30

(Simpson)

110 “

62

“ =56

less than 34 “

(hertwigi)

624

MEMOIRS OP THE CARNEGIE MUSEUM.

62. Anodontites riograndensis (Von Ihering) (1890).

Shells: Plate XLIII, figs. 2, 3; Plate XLTV, fig. 2.

Anodonta riograndensis A^on Ihering, 1890, pp. 154, 158.

Glabaris riograndensis Von Ihering, 1893, pp. 118, 119.

Anodonta exotica Corsi {non Lamarck), 1901, p. 456, fig. 37.

It is quite possible that A. exotica of D’Orbigny {non Lamarck) represents
chiefly this species. However, on account of the great diameter (44 pr. ct. of length)
of the specimen measured, this apjiears to be rather A. forbesiana. Since there is
no figure given, D’Orbigny ’s species remains doubtful. The figure of A. exotica
given by Corsi certainly in this s])ecies.

Type-loccditij. — Rio Grande do Sul, Brazil.

Other Localities.— Rio Paraguay (Von Ihering); Lower Parana and La Plata
Rivers (Von Ihering) ; Rio Uruguay (Von Ihering) ; De]jartment of Colonia, Uruguay
(Corsi); Arroyo Mendoza, Dejiartment Florida, Uruguay (Corsi); Montevideo,
Uruguay (Von Ihering).

New Localities. — In a i)ond along banks of the Rio Negro, Santa Isabel,
Uruguay (J. D. Haseman coll., Februaiy 11, 1909). Two males with soft parts,
four odd right vales. Headwaters of Rio Paraguay, Santa Rita, Matto Grosso,
Brazil (J. D. Haseman coll., June 12, 1909). One specimen. Rio de la Plata,
San Isidro, 20 km. N. of Buenos Aires, Argentina (A. Windhausen coll., January
1917). Three specimens, male, and gravid females, with soft parts.

Distribution. — La Plata-drainage from Buenos Aires up to the headwaters of
the Paraguay in Matto Grosso; in the Uruguay and its tributaries; and also in
streams of the Banda Oriental to Florida and Montevideo. According to Von
Ihering occurring also in the Rio Grande do Sul, but exact localities not known.

Characters of Shell. — Shell large (length up to 130 mm. and over), moderately
and variably solid, but never thin. Valves distinctly gaping anteriorly, slightly
so iiosteriorly. Outline subtraiiezoidal, rather elongated, height 54 to 63 pr. ct.
of the length, distinctly oblicpie. Ujiper margin gently curved or nearly straight,
forming a more or less distinct, obtuse angle with the posterior margin, and also
with the anterior margin. Sometimes in old siiecimens these angles are obliterated.
Posterior margin oblicpiely descending, not very steep, straight, or gently convex,
curving around into the jiosterior lower margin, forming a blunt posterior point
of the shell, situated a little above the base-line. Lower margin convex, but
ascending in its anterior jiart, and becoming here nearly straight for a distance just
in front of the middle; then curving up into the anterior margin. Thus the shell
is distinctly narrower anteriorly, and this produces the oblique shape. Often

ortmann: south American naiades.

625

this shape is obscured on an external view on account of the elevation of the beaks,
but it is best seen from the inside, when the ligament behind the beaks is placed
horizontally.

Valves moderately convex, somewhat flattened upon the sides, greatest diam-
eter 34 to 40 pr. ct. of the length, located upon the posterior ridge, some distance
behind the beaks. Posterior ridge indistinct. Posterior slope compressed, some-
times with traces of a radial ridge and furrow. Beaks a little inflated and a little
elevated above the hinge-line, their tips located at 29 to 33 pr. ct. of the length.

Epidermis rather smooth and shining. Irregular subconcentric ridges are
present, but no lamelliform striae, exce]it on the posterior slope,, where they may
be more or less distinct, and rather crowded. Upon the smooth part of the disk
there are often a few radial scalariform stripes, consisting of short subconcentric
wrinkles. But these may be entirely missing. Color of epidermis prevailingly
green, but shading into brown, often with concentric bands of darker or lighter
green and brown. Green rays may be present or absent; when present, they are
best developed on and just in front of the posterior ridge. Posterior slope generally
dark green to blackish, often with a few black rays.

Hinge-line nearly straight behind the beaks, but gently and distinctly curving
down anteriorly, so that the whole hinge-line appears as gently curved. A trace
of a slight sinuation may be observed in a very obscure elevation of the hinge-
line at its anterior end, but generally this is not the case. Ligamental sinus broad
and deep, in older specimens deeper than in younger ones, its anterior margin
running obliquely backward in young individuals, but being vertical in old ones.
Sometimes the anterior margin is curved, the lower point turning forwards.

Cavity of shell and beaks moderate, corresponding to the obesity of the shell.
Nacre whitish and iridescent, often much discolored, and quite frequently there
are fine, irregular, subconcentric, waved, or zig-zag, black lines inside of the shell,
chiefly near the pallial line. Faint radial striations may be present in young
specimens. Prismatic border narrow or very narrow, subequal in width, grayish
green or brownish green.

Anterior adductor-scar slightly impressed, irregular in outline, with an upper
process representing the anterior retractor-scar. Anterior protractor-scar separated
from adductor-scar. Posterior adductor-scar faint and often indistinct, the pos-
terior retractor forming an upper triangular process thereof. No dorsal scars.
Pallial line subconcentric to the margin.

Remarks. — ^The height is given by Von Ihering as from 49 to 57 pr. ct. of length,
thus being less on the average than in my specimens; but I think this is due to the

626

MEMOIRS OF THE CARNEGIE MUSEUM.

fact that Von Ihering did not measure in the way I did (See p. 526). If I measure
the length of my specimens along the longest axis (diagonally), and the height
vertically to it, I obtain for the above specimens figures more nearly agreeing with
those of Von Ihering from 51 to 59 pr. ct. of length.

Measurements.

LocaIitie.=5.

d

X

o

m

Length.

Height.

Diameter.

Beaks.

Figured.

Santa Isaljel .

3

?

95 nmi.

59.5niin

= 03 pr.ct. of L.^32 nun

= 34 pr. ct. of L.

at 28 nnn

= 29 pr.

ct. of L.

Do.

4

?

102

55 “

= 54

38 “

= 37

29.5 “

= 29

Do.

a

107

03

= 59

43 “

= 40

30 “

= 30

San Isidro. . .

y

9

123

74 “

= 00

48 “

= 39

30 “

= 29

PI. XLIII, f. 2.

Do.

X

cf

123.5 “

70.5 “

= 01

49 “

= 40

30 “

= 29

Pl.XLIII, f. 3.

Do.

Z

9

130

80 “

= 03

54 “

= 40

45 “

= 33

PI. XLIV, f. 2.

In other respects, the characters given b}" Von Ihering for riograndensis agree
with my specimens. This is especially true of the diameter, which, according
to Von Ihering ranges from 30 to 39 pr. ct., and in my specimens from 34 to 40 pr. ct.
Von Ihering believes that the diameter as well as the height differ according to
sex. According to my material there surely is no such differentiation. It is im-
possilile to directly comjmre my measurements for the location of the beaks (29
to 33 jir. ct.) with those given by Von Ihering, because he measured the distance
from the anterior end of the hinge-line (34 to 43 pr. ct.); but I should say that
measurements of my specimens taken in the same way give, for the above speci-
mens, the values: 36, 35, 37, 43, 40, 44 pr. ct. of length, agreeing fully with Von
Ihering’s.

The point of the beaks is inclined forwards, as Von Ihering mentions, but
not too much stress should be laid upon this character. An important feature,
however, is the curved hinge-line.

Glabaris trapesiolis cygneiformis Pilsbry (1896, p. 563, PL 26, figs. 4, 5) from
Maldonado, Uruguay, surely is closely allied. But, as Pilsbry points out, it is
more compressed than riograndensis (Diam. only 26 pr. ct.), and its posterior end
is more elevated above the liase-line; it is thus less oblique. For this reason I
cannot unite it with the present species, although it may fall under it.

Glabaris sinipsonianus Pilsbry (ibid., p. 564, PI. 27, fig. 13) from Rio de la
Plata, is also ver.y much like riograndensis. The height of 56 pr. ct. and the diam-
eter of 38 pr. ct. fall within the range of variation of riograndensis. But it is said
to be a very solid shell, of a rather regular, ol)long-oval shajie, with a large, elongated
anterior protractor scar. Having no specimens corresponding to it, I cannot express
an ojiinion.

Anatomy. — The soft ])arts of three males and two gravid females are at hand.

ortmann: south American naiades.

C27

the latter collected in January. Von Ihering found gravid females with lasidia on
May 28.

Anal opening entirely open, its inner edge smooth, separated from the branchial
opening by a mantle-connection. Branchial opening with fine papillae on inner
edge. Palpi very large, nearly semicircular, lower margins rounded, posteriorly
truncated, without a posterior point. Posterior margins widely separated at
base (this is a peculiar feature, not observed in any of the foregoing species).

Gills long and wide, the inner the wider anteriorly, its anterior end inserted
between the posterior ends of the palpi (this is also a peculiar feature). Outer
gill with the anterior end at the highest point of the mantle-attachment-line. Inner
lamina of inner gill entirely connected with abdominal sac. Gills with well de-
veloped septa running in the direction of the gill-filaments, alternately (but ir-
regularly so) stronger and weaker in the non-marsupial gills. The inner gill of
the female is marsupial, with stronger, more uniform septa, which, however, are
not more closely set, and have the usual vertical ridges near the outer lamina,
projecting into the lumen of the water-tubes. When charged, only the inner com-
partment is filled with ova, thus becoming an ovisac, while the outer compartment
remains a (secondary) water-tube. Egg-masses only loosely hanging together.
Eggs very small, according to Von Ihering 0.071 to 0.090 mm. in diameter, while
I have found them to be about 0.08 mm. In this species also I have not been able
to find mature larvae, but this is one of the species, in which Von Ihering has ob-
served the lasidium, which he describes as being 0.1 mm. long.

63. Anodontites forbesiana (Lea) (1860).

Shells: Plate XLIII, fig. 4; Plate XLIV, fig. 3.

Simpson, 1914, p. 1438.

Additional References. — Anodonta forbesiana Von Ihering, 1890, p. 158.
CoRSi, 1901, p. 457.

Glabaris forhesianus Pilsbry & Rush, 1896, p. 81.

Type-locality . — Uruguay River.

Other Localities. — Rio de la Plata, Colonia, Uruguay (Pilsbry & Rush).

New Localities. — Rio Uruguay (in mud), Uruguayana, Rio Grande do Sul,
Brazil (J. D. Haseman coll., February 5, 1909). Fourteen specimens, males and
females, all with soft parts. In pond along banks of Rio Negro, Santa Isabel,
Uruguay (J. D. Haseman coll., February 11, 1909). Two odd right valves.

Distribution. — Known only from the Rio de la Plata below the mouth of the
Uruguay, Rio Uruguay up to Rio Grande do Sul, and its tributary the Rio Negro.

628

MEMOIRS OF THE CARNEGIE MUSEUM.

Simpson also gives Peru, but I do not know on what authority, and strongly doubt
this record.

Von Ihering (1890) pointed out the differences from A. riograndensis, to
which this s])ecies is closely allied. The original specimens of A. Jorhesiana were
injured at the posterior end, and it was believed that in uninjured ones this end
might be more strongly produced. However, Lea’s figure shows in the growth-
lines that the shape of the shell, before it was injured, was similarly truncated, and
my series, which contains a majority of intact shells, demonstrates that the normal
shape of this s])ecies also exhibits this truncation, i.e., a steeply descending pos-
terior margin. This is the chief character of this species, and in consequence, the
dimensions are different from those of A. riograndensis.

This difference is seen first of all in the height, which varies from about 60 to
69 jir. ct., falling under 60 pr. ct. only in very young specimens, while in riograndensis
it is from 54 to 63 pr. ct. The diameter of the two species is about the same, but
slightly higher on the average in forbesiana (35 to 48 pr. ct. and in young ones
as low as 33 pr. ct. against 34 to 40 in riograndensis) . The beaks of forbesiana
are farther removed from the anterior end: 32 to 38 pr. ct., against 29 to 33
pr. ct. in riograndensis. Von Ihering gives the umbonal index as 48 pr. ct., but
this is due to his different method of measuring; measured in my specimens ac-
cording to his method, it would be from 42 to 56 pr. ct., while it is 34 to 43 pr. ct.
in riograndensis. The index above 50 ]ir. ct. is found only in my youngest speci-
mens, where the ligamental sinus is very anterior, thus making the hinge-line very
short.

Finally, in forbesiana, the hinge-line is practically straight, as mentioned by
Von Ihering, and represented in Lea’s figure. This holds good in all of my speci-
mens.

In othei’ respects this species agrees with riograndensis, but it should be re-
marked that the radial scalariform stripes are generally absent or very poorly
develojied; only in one or two cases a few of them are distinctly seen on the anterior
]iart of the shell.

Measurements (Specimens prom Uruouayana).

No.

Sex.

Length.

Heiglit.

Diameter.

Beaks.

Figured.

c. . .

c?(?)

29 mtii.

10 mm

= 55 pr.et. of L.

9.5mm

= 33 pr.ot. of L.

at 10 mm. =34 pr. ct. of L.

1. . .

&

49 “

29 “

= 59

17.5 “

= .30

17 “ =35

2.

cf

71 “

42.5 “

= 00

25 “

= .35

23 “ =32

4. . .

9

72 “

47 “

= 05

28 “

= 39

25.5 “ =35

7. . .

9

85 “

54.5 “

= 04

30 “

= 42

29 “ =34

8. . .

cf

87 “

59 “

= 08

42 “

= 48

33 “ =38

PI. XLIII, fig. 4.

10. . .

&

100 “

()9

= 09

43 “

= 43

37 “ =37

11. . .

9

108 “

70 “

= 05

40 “

= 43

38.5 “ =30

PI. XLIV, fig. 3.

Lea’s fig. . .

114 “

71

= 02

.54 “

= 47

44 “ =38

ortmann: south American naiades.

629

Simpson’s measurements do not agree with these figures.

It should be noticed that young specimens are not so high, and also less swollen
than older individuals, and thus the typical shape is less evident in them. They
are also extremel}' thin-shelled, while older ones are rather solid.

Anatomy. — I have nine males and five females. The structure is exactly like
that of A. riograndensis. The palpi have their posterior bases separated; for
about two-thirds of their length the bases are contiguous, but then they diverge.
The inner gill begins just without and close to the end of the inner palpus.

64. Anodontites rioplatensis (Sowerby) (1870).

Anodon rioplatensis Sowerby, XVH, 1870, PI. 26, fig. 101.

Glabaris trapesialis rioplatensis Simpson, 1900, p. 925.

Anodontites trapesialis rioplatensis Simpson, 1914, p. 1431.

Type-locality . — Rio de la Plata.

Other Localities. — Haas (1916, pp. 29, 54) mentions a closely related form from
Rio Uruguay, Salto Oriental, Uruguay, which may, or may not, be this.

New Locality. — Rio Limay, Patagonia, Argentina (received in exchange from
W. Israel). One specimen.

Distribution. — The Rio de la Plata and Rio Negro drainages in Argentina.
Possibly also in Rio Uruguay.

We may regard this as an exaggerated forbesiana, with the shell higher and
shorter, and the beaks more central, but in all other respects it is similar. Sowerby
calls the shell thin, and describes concentric “wrinkles” on the beaks. Our speci-
men is rather solid, and the wrinkles are nothing but growth-lines.

Measurements.

Length.

Height.

Diameter.

Beaks.

108 mm.

106 “

82 mm. = 76 pr. ct. of L.

76 “ =72

49.5 mm. =46 pr. c

t. of L.

at 4.3 mm. =40 pr. ct. of L.

(my speci-
men)

(Sowerby’s

figure)

Subgenus Lamproscapha Swainson (1840).

Lamproscapha Swainson, Treat, on Malacology, 1840, p. 381.

Virgula Simpson, 1900, p. 931; 1914, p. 1454.

Shell greatly elongated, knife-shaped, sharply pointed behind, with a sharp
posterior ridge. Posterior retractor-scar completely separated from the adductor-
scar, and remote from it by about one to three times its own diameter. Epidermis
not shining, covered with very fine concentric (posteriorly) and radial scalari-
form (anteriorly) wrinkles.

630

MEMOIRS OF THE CARNEGIE MUSEUM.

The greatly elongated shape and the posterior retractor-scar are evidently
correlated characters. This snbgeniis apparently stands close to the crispata-
groiiji of Anodontites in the sculpture of the ejiidermis.

Swainson introduced Lcmiproscapha for four species, of which the first {elongata
Swainson) was doubtfully referred here. The second is ensiformis. If the latter
is to be sejiarated from Anodontites, Lamproscapha is the oldest available name.

65. Anodontites (Lamproscapha) ensiformis (Spin) (1827).

Anodon ensiformis Spin & Wagner, 1827, p. 31, PI. 24, figs. 1, 2. Sowerby, XVII,
1867, PI. 11, fig. 31 (young).

Anodonta ensiformis D’Orbigny, 1843, p. 618, PI. 79, fig. 10. Von Ihering, 1890,

p. 161.

Glabaris ensiformis Simpson, 1900, p. 932.

Anodontites ensiformis Simpson, 1914, \). 1455. Haas, 1916, pp. 36, 57.
Type-locality not given.

Other Localities. — Rio San Miguel (= Rio Itonama), Bolivia (tributary to
Guapore) (D’Orbigny); Rio Piray, Santa Cruz de la Sierra, Bolivia (tributary to
Rio Marmore, into which the Guapore flows) (D’Orbigny); Rio Napo, Mazan,
Peru (Haas). Reiieatedly reiiorted from Brazil, but no exact localities given.

Lea (Obs. XHl, 1874, p. 27) mentions this sjiecies as having been found in
Guyana, in connection with a species from Yuruari River (tributary to Essequibo).
Simpson (1900, ]). 932 and 1914, p. 1456) describes a new species {A. falsa) taken
by Lea for ensiformis, from the same river (Yuruari), '‘a branch of the Orinoco.”
Tlie Yuruari, however, is a tributary of the Plsseiiuibo, but is located chiefly in
Venezuela, not in Guyana, and does not belong to the Orinoco-system.

New Locality. — Rio Machupo, San Joaquim, Bolivia (tributar}^ to Rio Itonama
and Guapore) (J. D. Haseman coll., September 5, 1909). Four specimens, three
of them with soft ]iarts. Another specimen is in the Carnegie Museum (from the
Hartman collection) labeled ^‘Brazil.”

Di.stributio'n. — Definite localities are so far known only from the iqiper Amazon
and Aladeira drainages in Peru and Bolivia.

Characters of the Shell. — Shell moderately thick; outline much elongated,
knife-like, ])ointed behind. Height 25 to 33 ])r. ct. of the length. Valves not
galling. Dorsal and ventral margins practically jiarallel, excejit towards the
])osterior end, where the dorsal margin curves in a gentle curve or a very obtuse
angle into the iiosterior margin, which descends obliquely and is straight or gently
concave. At the ])osterior end the margin curves sharply around to the ventral

ortmann: south American naiades.

631

margin, forming a blunt, but distinct point, which is hardly, or very little, elevated
above the base-line. Lower margin almost straight, l)ut with a more or less distinct
concavity in the middle. Anteriorly the lower margin curves up into the rounded
anterior margin.

Valves very slightly convex, practically flat upon the sides, and, in large
specimens, even with a shallow depression corresponding to the concavity of the
lower margin. The posterior ridge is distinct, but rounded, running towards the
posterior point of the shell. Above this ridge, the shell is somewhat compressed.
Diameter 14 to 20 pr. ct. of the length. Beaks low, and hardly elevated above
the hinge-line, located at 18 to 26 pr. ct. of the length, and proportionally more
anterior in older specimens; the large specimen, described by D’Orbigny, has
them at 15 pr. ct. of the length.

Epidermis not smooth, with irregular concentric lines, heaviest upon the pos-
terior ridge, and with very tine strise, sublamelliform upon the posterior slope and
towards the margins. Faint radiating lines are present, dividing the fine stria)
into scalariform stripes of crowded, fine wrinkles, visible only in well-preserved
specimens, and restricted to the anterior part of the shell. Color of epidermis
greenish or brownish olive, inclining to blackish in old shells, without color-mark-
ings and without distinct growth-rests.

Hinge-line straight behind the beaks, slightly tlescending in front of them.
Idgamental sinus much wider than deep, its anterior margin oblique to the hinge-
line, but more nearly vertical in the largest specimen at hand.

Cavity of shell and beaks very shallow. Nacre, in all of my specimens, whitish,
but with blueish and purplish iridescence (according to other authors, it is some-
times coppery), with irregular radiating lines, most distinct towards the margins.
Prismatic zone rather narrow, and subcoiicentric with the margin. Anterior ad-
ductor-scar well impressed, suliovate. Anterior retractor-scar distinct, but con-
nected with that of the adductor-scar. Anterior protractor-scar separated. Pos-
terior adductor-scar faint, subovate. Posterior retractor-scar separated from the
latter and rather removed from it, at least by as much as its own width ; in old shells
up to three times its width.

Remarks. — There can be no mistaking this species, the elongated shape being
so characteristic, that no other South American form could be taken for it, with
the possible exception of A. falsa (Simpson) from Venezuela (See above). It
constitutes with the latter a peculiar group within the genus, but approaches the
normal Anodontites (of the crispata-tjpe) more than any other. The location

632

MEMOIRS OF THE CARNEGIE MUSEUM.

of the posterior retractor-scar is quite unique, but, of course, is connected with
the elongated shape of the shell.

Measurements.

Locality.

No.

Length.

Height.

Diameter.

Beaks.

S. Joaqiiim

35 nim.

10 mm. =29 pr. ct. of L.

0 mm. = 17 pr. ct. of L.

at 9 mm. =26 pr. ct. of L.

Do

1

50 “

15 “ =30

10 “ =20

10 “ =20

Do

2

55 “

14 “ =25

9 “ =16

11 “ =20

Do

3

52 “

17 “ =33

10 " =19

10 “ =19

"Brazil ”

78 “

20 “ =26

12 " =15

14 “ =18

D'Orbignv

130 “

= 29

= 19

= 15

Simpson

07 “

25 “ =26

15 “ =15

Do

100 “

27 “ =25

15 “ =14

Anatomy. — Tlie soft parts of three specimens are at hand, but their sex is
not positively known.

The structure is that of the genus Anodontites, with such modifications as are
caused liy the elongation of the shell. The mantle-connection between anal and
branchial openings is a little longer than usual, the gills are extremely long and
narrow, and the part behind the foot, where the two inner laminae of the inner
gills are connected, is proportionally longer than in any other species. As indicated
by the scars of tlie shell, the posterior retractor muscle is considerably removed
from the adductor.

Anal opening entirely open. Branchial opening with very small papillae.
Palpi comparatively long, but narrow, lower margins forming a gentle curve. Pos-
teriorly they are very briefly truncated, forming the posterior margins, which
are not connected. Structure of gills as usual, with distinct septa. I have not
been able to positively identify the sex of the three specimens at hand: the con-
tlition of the gills is rather unsatisfactory, they being much torn, so that no sections
could be made, and from macroscopical examination (and with a lens) no indica-
tions of a differention of the inner gills could be detected : this, indeed, would indi-
cate the male sex, ljut my specimens are too young for one to be sure about this.
The inner gill, as usual begins immediately behind the palpi, and the inner lamina
of the inner gill is entirely connected with the abdominal sac. Foot rather long,
but altogether small; of course, the real shape could not be made out on account of
the contraction in alcohol.

Genus Mycetopoda D’Orbigny (1835).

Simpson, 1914, p. 1457.

Characterized by very elongated, siibtrapezoidal shell, which is widely gaping
in front. The chief characters, however, are in the soft parts. The foot is ex-

ortmann: south American naiades.

633

tremely elongated and dilated at the end (button-like), and probably is in life
never entirely withdrawn into the shell (hence the gaping margins). In addition,
the branchial opening is said to be closed below (Simpson), but I have not been
able to confirm this, and furthermore the pallial line does not show any indication
of this (having no sinus). D’Orbigny makes the positive statement that the bran-
chial is not closed. The anal opening is closed in part.

This genus has a wide distribution in South America, from the Cordilleras
eastward, and from Argentina northwards into Central America, at least as far
as Guatemala.

Von Ihering has given a key for the species (1910, p. 118); although this is
not always quite satisfactory, I have used it in the identification of my compara-
tively meagre material. Von Ihering’s treatment of the genus surely has cleared
up a good deal, but it is not to be regarded as final. His opinion that there are
species of this genus in Eastern Asia {Solcnaia) certainly is incorrect. We do not
know the anatomical structure of the latter, except that the foot is said to be
similarly developed. But we must not forget, that there is a North American
t/nfomd-shell {Lastena lata), which also has a foot like this. In the muscle-scars,
the ligamental sinus, and the beak-sculpture Solenaia undoubtedly differs from
the South American Mycetopoda, which according to the anatomy is a Muteline-
shell.

The species {subsinuata) of which 1 have studied the anatomy, has another
character, the partly closed anal-opening, in which it differs from all South American
Mutelince, and resembles the African members of this subfamily. This may be
another peculiar feature of the genus, but it is desirable that other species should
be examined as to this.

66. Mycetopoda siliquosa (Spin) (1827).

Mycetopoda siliquosa Von Ihering, 1910, p. 120; Simpson, 1914, p. 1458.
Mycetopoda bahia Von Ihering, 1910, p. 122, PI. 12, fig. 3; Simpson, 1914, p. 1463.

According to Simpson, M. legumen (Von Martens) and M. clessini Von Ihering
belong here.

Type-locality. — Rio Paraguassii, Bahia, Brazil.

Other Localities.— Rio Piracicaba, Piracicaba, Sao Paulo, Brazil (Von Ihering,
1893); Rio Sao Francisco, Villa Nova, Sergipe (not Bahia), Brazil (Von Ihering,
M. bahia).

New Locality. — Lagoa Salgado, Bahia, Brazil (upper Rio Salitre, tributary to
Sao Francisco) (J. D. Haseman coll., November 10, 1907). Two specimens. Two

634

MEMOIRS OF THE CARNEGIE MUSEUM.

other siiecimens are in the Carnegie Museum, without exact localities, from the
Holland and Juny collection resi)ectivel30

Distribution. — Restricted to eastern Brazil, to the drainages of the Rio Parana
in Sao Paulo, and of the Rio Paraguassu and Rio Sao Francisco in Bahia and
Sergipe. Possibly more widely distributed in the Sao Francisco system.

Haas (1916, p. 37, 58) gives this species also from Rio Unuyacu (tributary
to Rio Napo) in Ecuador; however, he conceived it in Simpson’s sense, and we
cannot be sure that it is M. siliquosa as defined by Von Ihering.

My specimens fully agree with the account given by Von Ihering, and their
measurements come very close to those given by him.

Measurements.

Locality.

Length.

Height.

Diameter.

Beaks.

Lagoa Salgado

72 mill.

26

mill. =36 pr. ct. of L.

13.5

mm. = 17 pr. ct. of L.

at 19 mm

= 26 pr. ct. of L.

Holland coll

80

29..'

“ =.37

15

“ =19

20 “

= 25

.lunv coll

85.5 “

31

“ =36

15

“ =18

21 “

= 25

Lagoa Salgado

89

35

“ =39

20

“ =22 “

27 “

=30

Spix’ type

80

31

“ =.39

22 “

= 28

The figures for Spix’ type are taken from Von Ihering (1890). Von Ihering
(1910, in the key) gives the location of the beaks as ranging from 18 to 29 pr. ct.
and the height as ranging from 35 to 37 pr. ct. of the length.

Remarks. — In all of my specimens, the posterior adductor scar is presinual.
ill. bahia is founded upon a single specimen, which has the measurements: L. 78,
H. 27 = 35 pr. ct., F). 15.5 = 20 pr. ct., beaks at 23 = 30 pr. ct. These figures
fall within the range of variation of M. siliquosa. In Von Ihering’s key there is
here a weak point, since the forms are distinguished chiefly by the location of the
beaks, with the figures partly overlapping. I cannot find any difference in bahia
from siliquosa, except that the lower margin in the former ascends slightly behind,
and that the iiosterior adductor scar is said to be '‘subsinual, in part even a little
presinual.” The first character very well may be individual; the second, dis-
regarding th(? fact that it is hard to understand, does not at all differ from siliquosa,
where this scar is simply presinual. Indeed, my young specimen from Lagoa
Salgado has the adductor-scar less in advance of the ligamental sinus than the
larger. Altogetlu'r, this young specimen is extremely close to bahia, only the
posterior lower margin is not curved up, and the beaks are more anterior. It is
also remarkable for the great thinness and transparency of the shell. Therefore
I think that M. bahia is only a young individual of M. siliquosa.

oetmann: south American naiades.

635

67. Mycetopoda staudingeri (Von Ihering) (1890).

Mycetopus staudingeri Von Ihering, 1890, p. 130, figs. A, B.

Mycetopoda staudingeri Simpson, 1900, p. 934; Von Ihering, 1910, p. 121.
Mycetopoda siliquosa staudingeri Simpson, 1914, p. 1460.

Type-locality. — Rio Hiiayabamba, Peru (tributary to Rio Huallaga).

Additional Locality. — Rio Huallaga, Peru (Von Ihering).

Locality Represented in Carnegie Aluseum. — Marafion, Upper Amazon (Hart-
man collection). One specimen.

Distribution. — Headwaters of the Amazon in Peru; a variety {cequatorialis
Von Ihering) in Ecuador.

Our specimen agrees very well with Von Ihering’s description and figures,
chiefly with fig. A (which is supposed to be a male) but it to a certain degree stands
between figs. A and B (See measurements of height). The postsinual position
of the posterior adductor-scar is evident and remarkable, and it seems, indeed,
that this is an important taxonomic character. The posterior end of the upper
margin (behind the ligamental sinus) is in my specimen not so greatly elevated
as in Von Ihering’s figures.

Measurements.

Length.

Height.

Diameter.

Beaks.

My specimen

94 nim.

.38 mm. =40 pr. ot. of L.

19 mm. =20 pr. ct. of L.

at 24 mm. =25 pr. ct. of L.

Von Ihering, A

9.3 “

35 “ =.38

Do. \ B

103 “

43 “ =12

21 “ =20

In 1910, Von Ihering gives for the height 35 to 37 pr. ct. of the length, which
does not exactly agree with his original figures. The location of the beaks is ac-
cording to him is 27 to 28 pr. ct. of the length.

68. Mycetopoda subsinuata (Sowerby) (1868).

Mycetopus subsinuatus Sowerby, XVI, 1868, PI. 4, fig. 10; Von Martens, 1900,
p. 540, PI. 41, fig. 5.

Mycetopoda subsinuata Simpson, 1900, p. 934; Von Ihering, 1910, p. 120; Simpson,
1914, p. 1461.

Type-locality . — Bogota, Colombia.

Other Localities. — Ecuador (Von Ihering) ; Paso Antonio, West Guatemala
(Pacific slope) (Von Martens).

New Localities. — Maranon, Upper Amazon (Peru) (Hartman coll.). Three
complete specimens, four left valves. Rio Conchins, Maya Farm, Quirigua,
Guatemala (Atlantic slope, to Rio Montagua) (A. A. Hinkley coll., February 6,

636

MEMOIRS OF THE CARNEGIE MUSEUM.

1913). One shell, female, with soft parts, and soft parts of a male and female with-
out shells.

Distribution. — From the upper Amazon drainage in Peru and Ecuador through
Colombia into Central America, northward to Guatemala, where it is found both
on the Atlantic and Pacific slopes.

Von Ihering’s key has failed me in the identification of this species, since he
uses size as a criterion, while my specimens are all rather small. The height,
also used as a distinctive character, is apparently unreliable, as shown by my
specimens. Therefore this grouj) {hh in the key) needs revision.

Nevertheless the two largest specimens at hand (one from the upper Amazon,
the other from Guatemala) agree fairly well with Von Ihering’s account and
measurements, and also with those of Von Martens. There is no question about
the identity of the Central American specimens with the form from northern South
America. The posterior adductor-scar is said to be subsinual (Von Ihering p. 118).
This fits my smaller specimens from the upper Amazon, while the larger one is
lieculiar. It apiiears as if there were in each valve two superimposed scars, the
one more anterior, the other a little farther back. The latter is superficial, but ap-
parently corresponds to the latest growth-addition to the shell. Its anterior end
is slightly in advance of the ligamental sinus (iiresinual) . In this specimen, how-
ever, there is a disturbance of the regular growth, as indicated by a strong growth-
rest on the outside of the shell, and the shell looks as if stunted behind. This
individual is therefore not normal. In the large specimen from Guatemala, the
scar is also slightly presinual (about one-third of the retractor-process projecting
in front of the sinus), and in Von Vlartens’ figure (representing a very large speci-
men) this scar is still more presinual.

The sinuosity of the lower margin is seen in my two larger specimens, but is
not so strong as in Sowerby’s and Von Martens’ figures. The young specimens,
which undoubtedly belong with the larger one, show hardly a trace of this sinuosity.

Measurements.

Locality.

No.

Sex.

Length.

Height.

Diameter.

Beaks.

Maranoii

1

V

58

nun.

17 mm. =29 pr. ct. of L.

10

mm. =17 pr. ct. of L.

at 15

mm. =26 pr. ct. of L.

Do

2

'}

63

“

19 “ =30

10

“ =16

17

“ =27

Do

3

75

23 “ =31

12

“ =16

20

“ =27

Do

4

V

75

“

23 “ =31

12

“ =16

20

“ =27

Guatemala

9

S6

28 “ =33

17.5

" =20

21.5

“ =25

Maranon

5

(88)

“

31 “ =(35 “ )

20

“ =(23 “ ■)

26

“ =(30 “ )

So\verl>v

118

“

41 “ =35

Von Martens

125

“

43 “ =34

23

“ =18

Do

132

45 “ =34

23

“ =17

29

“ =32

ortmann: south American naiades.

637

My specimen, No. 5, from the Marahon is the one, which probably is injured, and
the length probably would be greater, when normal; this, of course, has influence on
the other indices, which are all somewhat too high. The figures taken from Sowerby
and Von Martens are close to mine. The greater height undoubtedly is due to
the larger size of the specimens, and probably also to the more anterior location of
the beaks.

Anatomy. — Soft parts of one male and two females at hand, one of the latter
is gravid (collected February 6).

The anatomy is that of the South American Mutelinoe, but it resembles that
of the African Mutelince, in having the anal opening closed above for about half
of its length. The opening is therefore comparatively short, being only a little
longer than the branchial opening, and reaching upward only to about the middle
of the adductor muscles. In other South American MutelincE it reaches upward
beyond the posterior retractor-muscles. The anal is sejiarated from the branchial
opening by a connection of the mantle; its inner edge is smooth. Branchial opening
with fine papillae on the inner edge. Although Simpson says that the branchial
opening is closed below, I cannot find any trace of a mantle-connection at the lower
(anterior) end. There is also no indication that such a connection has been torn
during life, or in preservation. Palpi long and low, their lower margins curved,
not drawn out into a posterior point, posteriorly with a short truncation, forming
the posterior margins, which are not connected.

Foot very large, subcylindrical and subcompressed, at the distal end swollen
into a button-like knob. This structure is likewise not seen in other Muteline
shells.

Gills long and narrow, the inner the wider, the anterior ends as usual. Inner
lamina of inner gill entirely connected with abdominal sac. Distinct, continuous
septa of the Muteline type are present. The inner gill of the female is inarsupial,
with thicker, but not more crowded, septa; the water-tubes are again divided by
vertical ridges into an inner and outer compartment, the inner of which serves as
ovisac, the outer as secondary water-canal. Eggs small, loosely hanging together.
I have not been able to find mature larvas in my gravid female.

Genus Leila Gray (1840).

Simpson, 1914, p. 1399.

The chief character of this genus is the sinus of the pallial line below the pos-
terior adductor-scar, which is said to be connected with the closing of the branchial
opening at its lower (anterior) end, but particulars about this are not known.

In other respects the shell of this genus is very similar to that of the species

638

MEMOIRS OP THE CARNEGIE MUSEUM.

of the trapesialis-group of A7iodontites, with which it also has in common the gaping
margins. A peculiar character, generally missing in South American Mutelince,
is the jiresence of an olilique row of dorsal muscle-scars in the beak-cavity.

There can be hardly any question that this genus represents a more highly
siiecialized type of the genus Aiiodontites, and that its root is in the trapesialis-
group. Its distribution extends over South America, east of the Andes, and from
the liasin of the Amazon southward to northern Argentina.

The genus Leila is in great confusion, although there seem to exist only a
few species. Its revision has been attempted by Von Ihering (1890, p. 39) and
by Simpson, (1900, p. 914), but these two authors have arrived at very different
conclusions. Both agree in recognizing two groups: the one containing species
with a practically straight hinge-line (L. hlainvilleana and L. spixi), the other
species with a curved or sinuate hinge-line. While Simpson unites all of the forms
belonging to the latter group, into one species (L. esula), Von Ihering distinguishes
three as valid.

My material is entirely insufficient to iiermit me to decide this point. The
specimens before me unquestionaly come under L. castelnaudi Hupe, and so I shall
record them under this name, without attempting to pass upon the question
whether then are different from esula D’Orbigny and from pulvinata Hupe. They
also agree fairly well with Von Ihering’s descriiition of castelnaudi.

69. Leila castelnaudi Hupe (1857).

Leila castelnaudi Hupe, 1857, p. 91, PI. 19, fig. 1.

Anodon castelnaudi Sowerby, XVII, 1868, PL 20, fig. 79.

Columba castelnaudi Von Ihering, 1890, pp. 139, 142.

According to Simpson (1914, p. 1401) this is identical with L. esula (D’Orbigny).

Type-locality. — “Bourbon on Olympo, Paraguay.” This probably is Fuerte
Olympo, on Rio Paraguay, northern Paraguay.

New Localities. — Rio I^araguay, Corumba, Matto Grosso, Brazil (H. H.
Smith coll.). One left valve. Swamp of Lambare, Asuncion, Paraguay (J. D.
Hasenian coll., March 31, 1909). One left valve.

]\Ieasukement.s.

Locality.

I.ength.

Hciglit.

Diameter.

Beaks.

Hinge-line.

V. IhcriiiK

13S mill.

90 null. =70 pr.oi. of L.

105

mm. =76 pr.ct. of L.

Aisuiicion .

158

112 “ =71

88 mm. =43 pr. ct. of L. at 48 mm. =30 pr. ct. of L.

101

“ =64

CorunibA .

160 “

116 “ =70

70 “ =42

52 “ =31

114

“ =69

Von Ihering has measured according to a peculiar method, but I see no par-

ortmann: south American naiades.

639

ticular advantage in this. Yet the length of the hinge-line from its anterior end
to the anterior end of the ligamental sinus, and its proportion to the length of the
shell, should be noted, because there are slight differences in our specimens in this
respect.

According to Hupe’s figure, the beaks are located at 44 mm. = 32 pr. ct. of
length. This and the height given by Von Ihering agree very well with my speci-
mens, but the proportion of the hinge-line to the total length is less in the latter.
But the figures for my specimens show that that there is variation in this respect.

-BIBLIOGRAPHY.

1858 Adams, H. & Adams, A. The Genera of Recent Mollusca. Vol. II, London,
1858.

1906 Bartsch, P. Descriptions of Two New Naiades (Proc. U. S. Nat. Mus., XXX,
1906, pp. 393-395).

1900-1901 CoRSi, A. F. Moluscos de la Republica Oriental del Uruguay (Anal. Mus.
Nac. Montevideo, II, 1900, pp. 445-448; 1901, pp. 449-481).

Eigenmann, C. H. The Localities at which Mr. John D. Haseman Made Col-
lections (Ann. Carnegie Mus. VII, 1911, pp. 299-314).

Frierson, L. S. A New Pearly Freshwater Mussel of the Genus Hyria from
Brazil (Proc. U. S. Nat. Mus., XLVII, 1915, p. 363).

Germain, L. Mission du Service geographiqiie de I’Armee pour la Mesure d’un
Arc de Meridien Equatorial en Amerique du Slid. IX, Zool., Fasc. 3, Etude
sur les Mollusques Terrestres et Fluviatiles. 1910, pp. C. 63-C. 68.

Griffith, E. The Animal Kingdom by the Baron Cuvier. XII, 1834.

Haas, F. Nayades del Viaje al Pacifico (Trab. Mus. Cienc. Nat. Madrid.,
Zool., No. 25, 1916, pp. 1-63).

Haseman, J. D. A brief Report upon the Expedition of the Carnegie Museum
to Central South America (Ann. Carnegie Mus., VII, 1911, pp. 287-299).
Hupe, H. Mollusques, in Castelnau, F. De, Animaux nouveaux ou rares
recueillis pendant I’Expedition dans les parties centrales de I’Amerique du
Sud. 1857, pp. 1-103.

Ihering, H. Von, Revision der von Spix in Brasilien gesammelten Najaden
(Arch. f. Naturgesch., 1890, I, pp. 117-170).

1891-1892 Ihering, H. Von, Anodonta und Glabaris (Zoolog. Anzeig., XIV, 1891,
pp. 474-484; Ibid., XV, 1892, pp. 1-5).

1893 Ihering, H. Von, Najaden von S. Paulo und die geographische Verbreitung der
Suesswasserfauna von Siidamerika (Arch. f. Naturgesch., 1893, I, pp. 45-
140).

1910 Ihering, H. Von. Ueber Brasilianische Najaden (Abh. Senckenberg. Naturf.
Ges., XXXII, 1910, pp. 113-140).

1911

1915

1910

1834

1916

1911
1857

1890

640

MEMOIRS OP THE CARNEGIE MUSEUM.

1915 Ihering, it. Von. Molluscos, in Commissao cle Linhas Telegraphicas Estra-
tegicos de Matto Grosso ao Amazonas. Ann. V, 1915, pp. 1-14.

1819 Lamarck, J. B. de. Histoire Naturelle des Aniinaiix sans Vertebres, VI, 1819.
1832-1874 Lea, I. Observations on the Geiiiis LTnio. Vols. I-XIII, 1832-1874.

1870 Lea, I. Synopsis of the Family Unionidse. Fourth edition, 1870.

1917 Marshall, W. B. New and Little-known Species of South American Fresh-

water Vlussels of the Genus Diplodon (Proc. U. S. Nat. Mus. LIII, 1917, pp.
381-388).

1885 Martens, E. Von. Ueber brasilische Land- und Suesswasser-Mollusken. (Sitz.
Ber. Gesellsch. naturf. Fr. Berlin, 1885, pp. 147-149).

1894 Martens, E. Von. Die von Dr. Bohls in Paraguay gesammelten Mollusken

(Sitz. Ber. Gesellsch. naturf. Fr. Berlin, 1894, pp. 163-170).

1895 Martens, E. Von. Mollusken von Paraguay (Sitz. Ber. Gesellsch. naturf.

Fr. Berlin, 1895, pp. 33-35).

1900 Martens, E. Von. Biologia Centrali-Americana. Land and Freshwater Mol-
lusca. 1900-1901. (Unionidie, pp. 478-540, 1900).

1893 Nehring, a. Ueber Najaden von Piracicaba in Brasilien (Sitz. Ber. Gesellsch.
naturf. Fr. Berlin, 1893, pp. 159-167).

1918 Odhner, N. H. Zur Kenntnis der Homologien des Bivalvenschlosses (Geolog.

Foren. Stockholm Forhandh, 1918, pp. 562-590).

1835 Orbigny, a. D’. Synopsis terrestrium et fluviatilium Molluscorum in suo per
Americam Meridionalem Itinere (Magas, de Zoolog., 1835).

1843 Orbigny, A. D’. Voyage dans I’Amerique Meridionale, V, part 3, Mollusques.
1835-1843.

1910 Ortmann, a. E. The Soft Parts of Spatha kamerunensis Walker (Nautilus,
XXIV, 1910, pp. 39-42).

1911a Ortmann, A. E. The iVnatomical Structure of Certain Exotic Naiades compared
with that of the North American Forms (Nautilus, XXIV, 1911, pp. 103-108;
114-120; 127-131).

19116 Ortmann, A. E. A Monograph of the Naiades of Pennsylvania (Mem. Car-
negie Mus., IV, 1911, pp. 279-347).

1911c Ortmann, A. E. The use of the Generic Names Unio, Margaritana, Lymnium,
and Elliptic, and of Anodonta and Anodontites (Nautilus, XXV, 1911, pp.
88-191).

1912 Ortmann, A. E. The Anatomy of the Naiad Hyridella australis (Lamarck)
(Nautilus, XXV, 1912, pp. 100-103).

1918 Ortmann, A. E. The Anatomy of two African Naiades, Unio caller and Spatha
wahlbergi (Nautilus, XXXI, 1918, pp. 75-78).

1893 PiLSBRY, H. A. Notes on the Genera of Unionidae and Mutelidae (Nautilus,
Vn, 1893, p. 30).

ortmann: south American naiades.

641

1896 PiLSBRY, H. A. New Species of Freshwater Mollusks from South America (Proc.

Acad. Nat. Sci. Philadelphia, XLVIII, 1896, pp. 561-565).

1911 Pilsbry, H. a. Non-marine Mollusca of Patagonia, in Rep. Princeton Univ.

Exped. to Patagonia, III, part 5, 1911, pp. 609-610.

1896 Pilsbry, H. A. & Rush, W. H. List, with Notes, of Land and Freshwater Shells
Collected by Dr. Wm. H. Rush in Uruguay and Argentina (Nautilus X,
1896, pp. 76-79).

1864-1870 Reeve, L. A. & Sowerby, G. B. Conchologia Iconica. Vols. XVI-XVII,
(genera Unio, Mycetopus, Anodon, Hyria, Castalia) 1864-1870.

Rush, W. H., see: Pilsbry & Rush.

1900 Simpson, C. T. Synopsis of the Naiades or Pearly Freshwater Mussels (Proc.

U. S. Nat. Mus., XXII, 1900, pp. 501-1044).

1914 Simpson, C. T. A Descriptive Catalogue of the Naiades or Pearly Freshwater
Mussels. Detroit, 1914.

Sowerby, G. S., see Reeve & Sowerby.

1827 Spin, J. B. be & Wagner, J. A. Testacea fluviatilia quse in itinere per Brasiliam
collegit. 1827.

1840 SwAiNSON, W. A Treatise on Malacology. London., 1840.

Wagner, J. A., see Spin & Wagner.

(Note: As to the authorship of the species described by Spix and Wagner, Cf. Von
Ihering, 1890, pp. 118-119, footnote).

642

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXXIV.

Shells of Diplodon hasemani Ortmann and Diplodon imitator Ortmann.

All figures natural size.

Figs. 1-4. Diplodon hasemani Ortmann, from Rio Guapore, near Rio Sao Simao,
Matto Grosso, Brazil. Cam. Mus. Cat. No. 61.5857.

Fig. la. Adult male (No. 10), lateral view.

Fig. 15. Adult male (No. 10), dorsal view.

Fig. 2a. Adult, gravid female (No. 9), lateral view.

Fig. 25. Adult, gravid female (No. 9), inner view of left valve.

Fig. 2c. Adult, gravid female (No. 9), inner view of right valve.

Fig. 3. Half-grown, gravid female (No. 4), lateral view.

Fig 4. Young, gravid female (No. x), lateral view.

Figs. 5-7. Diplodon imitator Ortmann, from Rio Vaccahy-mirim, Santa Maria,
Rio Grande do Sul, Brazil. Cam. Mus. Cat. No. 61.9248. (See also PI. XXXV, figs. 1
and 2) .

Fig. 5a. Half-grown male (No. 29), lateral view.

Fig. 55. Plalf-grown male (No. 29), inner view of left valve. ‘

Fig. 5c. Half-grown male (No. 29), inner view of right valve.

Fig. 5d. Half-grown male (No. 29), dorsal view.

Fig. 6a. Young male (No. 14), lateral view.

Fig. 65. Young male (No. 14), dorsal view.

Fig. 7a. Adult female (No. 33), inner view of left valve (see also PI. XXXV, fig. 1).
Fig. 75. Adult female (No. 33), inner view of right valve.

Memoirs Carnegie Museum, Vol. VIII.

Plate XXXIV.

Diplodon.

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644

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXXV.

Shells of Diplodon imitator Ortmann, Diplodon simillimus Ortmann, and

Diplodon vicarius Ortmann.

All figures natural size.

Figs. 1 and 2. Diplodon imitator Ortmann, from Rio Vaccahj^-mirim, Santa Maria,
Rio Grande do Sul, Brazil. Cam. Mus. Cat. No. 61.9248 (See also PI. XXXIV, figs. 5-7).
Fig. 1. Adult female (No. 33), lateral view (same specimen as that figured on

PI. XXXIV, fig. 7).

Fig. 2. A"oung, gravid female (No. 21), lateral view.

Figs. 3-6. Diplodon simillimus Ortmann, from Rio Nhundiaquara, Morretes,
Parana, Brazil. Cam. Mus. Cat. No. 61.9250.

Fig. 3. Half-grown male (No. 11), lateral view.

Fig. 4. Half-grown male (No. 16), lateral view.

Fig. 5a. Adult, gravid female (No. 22), lateral view.

Fig. 55. Adult, gravid female (No. 22), dorsal view.

Fig. 5c. Adult, gravid female (No. 22), inner view of left valve.

Fig. 5d. Adult, gravid female (No. 22), inner view of right valve.

Fig. 6a. A'oung, gravid female (No. 24), lateral view.

Fig. 66. Young, gravid female (No. 24), dorsal view.

Figs. 7 and 8. Diplodon vicarius Ortmann, from creek (Rio Ribeira drainage).
Aqua Quente, near Iporanga, Sao Paulo, Brazil. Cam. Mus. Cat. No. 61.9251 (See also
PI. XXXVI, figs. 1 and 2).

Fig. 7. Half-grown male (No. 13), lateral view.

Fig. 8. Adult female (No. 15), lateral view (See also PL XXXVI, fig. 1).

Memoirs Carnegie Museum, Vol. VIII.

Plate XXXV.

Diplodon.

646

MEMOmS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXXVI.

Shells of Diplodon vicarius Ortmann, Diplodon decipiens Ortmann, and

Diplodon hild.e Ortmann.

All figures natural size.

Figs. 1 and 2. Diplodon vicarius Ortmann, from creek (Rio Ribeira drainage),
Aqua (Fiente, near Iporanga, Sao Paulo, Brazil. Cam. Mus. Cat. No. 61.9251 (See
also PI. XXXV, figs. 7 and 8).

Fig. la. Adult female (No. 15), dorsal view (same specimen as that figured on

PI. XXXV, fig. 8).

Fig. 16. Adult female (No. 15), inner view of left valve.

Fig. Ic. Adult female (No. 15), inner view of right valve.

Fig. 2. Half-grown, gravid female (No. 9), lateral view.

Figs. 3-6. Diplodon decipiens Ortmann, from creek (tributary to Rio Iguassii),
Serrinha, Parana, Brazil. Cam. Mus. Cat. No. 61.9253.

Fig. 3a. Adult male (No. 4), lateral view.

Fig. 36. Adidt male (No. 4), inner view of left valve.

Fig. 3c. Adult male (No. 4), inner view of right valve.

Fig. 3d. Adult male (No. 4), dorsal view.

Fig. 4. Adult female (No. 6), lateral view.

Fig. 5. Half-grown, gravid female (No. 3), lateral view.

Fig. 6a. Young female (No. c), lateral view.

Fig. 66. Young female (No. c), dorsal view.

Fig. 7. Diplodon hildee Ortmann, from Rio Jacuhy, Cachoeira, Rio Grande do
Sul, Brazil. Cam. Mus. Cat. No. 61.5864 (See also PI. XXXVII, figs. 1-3).

Fig. 7a. Nearly full grown male (No. q), lateral view.

Fig. 76. Nearly full grown male (No. q), dorsal view.

Memoirs Carnegie Museum, Vol, VIII.

Plate XXXVI,

Diplodon.

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648

MEMOIRS OP THE CARNEGIE MUSEUM,

EXPLANATION OF PLATE XXXVII.

Shells of Diplodon hild^ Ortmann and Diplodon mogymirim Ortmann.

All figures natural size.

Figs. 1-3. Diplodon hildce Ortmann, from Rio Jacuhy, Cachoeira, Rio Grande do
Sul, Brazil. Cam. AIus. Cat. No. 61.5864 (See also PI. XXXVI, fig. 7).

Fig. la. Young male (No. d), lateral view.

Fig. 16. Young male (No. d), dorsal view.

Fig. 2a. Adult female (No. 15), lateral view.

Fig. 26. Adult female (No. 15), dorsal view.

Fig. 2c. Adult female (No. 15), inner view of left valve.

Fig. 2d. AduT female (No. 15), inner view of right valve.

Fig. 3. Half-grown female (No. h), lateral view.

Figs. 4-7. Diplodon mogymirim Ortmann, from creek (tributary to Rio Mogy
Guassii), Mogy Mirim, Sao Paulo, Brazil. Cam. Mus. Cat. No. 61.9260.

Fig. 4a. Adult male (No. 22), lateral view.

Fig. 46. Adult male (No. 22), inner view of left valve.

Fig. 4c. Adult male (No. 22), inner view of right valve.

Fig. 5a. Half-grown male (No. 9), lateral view.

Fig. 56. Half-grown male (No. 9), dorsal view.

Fig. 6a. Young male (No. 12), lateral view.

Fig. 66. Young male (No. 12), dorsal view.

Fig. 7a. Adult female (No. 38), lateral view.

Fig. 76. Adult female (No. 38), dorsal view.

Memoirs Carnegie Museum, Vol, VIII.

Plate XXXVII.

Diplodon.

650

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXXVIII.

Shells of Diplodon berths Ortmann and Diplodon enno Ortmann.

All figures natural size.

Figs. 1-4. Diplodon berthce Ortmann, from Rio Jacuhy, Cachoeira, Rio Grande
do Sul, Brazil. Cam. Mus. Cat. No. 61.5865.

Fig. la. Adult male (No. 24), lateral view.

Fig. Ih. Adult male (No. 24), dorsal view.

Fig. Ic. Adult male (No. 24), inner view of left valve.

Fig. Id. Adult male (No. 24), inner view of right valve.

Fig. 2a. Young male (No. 3), lateral view.

Fig. 2h. Young male (No. 3), dorsal view.

Fig. 3a. Gravid female (No. 10), lateral view.

Fig. 36. Gravid female (No. 10), dorsal view.

Fig. 4. Gravid female (No. 16), lateral view.

Figs. 5-8. Diplodon enno Ortmann, from Rio Grande (Sao Francisco drainage),
Boqueirao, Bahia, Brazil. Cam. Mus. Cat. No. 61.9264.

Fig. 5. Half-grown male (No. 4), lateral view.

Fig. 6a. Young male (No. 12), lateral view.

Fig. 66. AYung male (No. 12), dorsal view.

Fig. 7. Adult female (No. 1), lateral view.

Fig. 8a. Nearly adult female (No. 2), lateral view.

Fig. 86. Nearly adult female (No. 2), dorsal view.

Fig. 8c. Nearly adult female (No. 2), inner view of left valve.

Fig. 8d. Nearly adult female (No. 2), inner view of right valve.

Memoirs Carnegie Museum, Vol. VIII.

Plate XXXVIII.

Diplodon.

652

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XXXIX.

Shells of Diplodon deceptus (Simpson), Diplodon (Cyclomya) paranensis (Lea),

AND Castalia undosa Von Martens.

All figures natural size.

Figs. 1-5. Diplodon deceptus (Simi^son), from Rio Jacuhy, Cachoeira, Rio Grande
do Sul, Brazil. Carn. Mus. Cat. No. 61.5868.

Fig. la. Adult male (No. 10), lateral view.

Fig. 16. Adult male (No. 10), dorsal view.

Fig. Ic. Adult male (No. 10), inner view of left valve.

Fig. Id. Adult male (No. 10), inner view of right valve.

Fig. 2. Adult male (No. 11), lateral view.

Fig. 3. Half-grown male (No. 4), lateral view.

Fig. 4. Half-grown female (No. 6), lateral view.

Fig. 5a. Young specimen (sex ?) (No. 2), lateral view.

Fig. 56. Young specimen (sex ?) (No. 2), dorsal view.

Figs. 6 and 7. Diplodon (Cyclomya) paranensis (Lea).

Fig. 6. Nearly adult female (No. a), lateral view, from Rio de la Plata, San Isidro,
Argentina. Carn. Mus. Cat. No. 61.9265.

Fig. 7. Young specimen, lateral view, from Rio Paraguay, Corumba, Matto
Grosso, Brazil. Carn. Mus. Cat. No. 61.2031.

Fig. 8. Castalia undosa Von Martens, from Rio Tiete, Itapura, Sao Paulo, Brazil.
Carn. Mus. C’at. No. 61.5116.

Fig. 8a. Half-grown specimen, lateral view.

Fig. 86. Half-grown specimen, dorsal view.

Fig. 8c. Half-grown specimen, inner view of left valve.

Fig. 8d. Half-grown specimen, inner view of right valve.

Memoirs Carnegie Museum, Vol. VIII.

Plate XXXIX.

Diplodon and Castalia.

654

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XL.

Shells of Prisodon alatus (Sowerby), Monocondyl.ea obesa Ortmann, and

Anodontites crispata Bruguiere.

All figures natural size.

Figs. 1-3. Prisodon alatus (Sowerby), from Rio Tapajos, Santarem, Para, Brazil.
Cam. AIus. Cat. No. 61.5109.

Fig. 1. Adult specimen, lateral view.

Fig. 2a. Half-grown specimen, lateral view.

Fig. 25. Half-grown specimen, dorsal view.

Fig. 2c. Half-grown specimen, inner view of left valve.

Fig. 2d. Half-grown specimen, inner view of right valve.

Fig. 3. Young specimen, lateral view.

Figs. 4-6. Monocondylcca ohesa Ortmann, from Rio Tapajos, Santarem, Para,
Brazil. Cam. Mus. Cat. No. 61.5850.

Fig. 4a. Half-grown specimen (No. 10), lateral view.

Fig. 45. Flalf-grown specimen (No. 10), inner view of left valve.

Fig. 4c. Half-grown specimen (No. 10), inner view of right valve.

Fig. 5a. Young specimen (No. 5), lateral view.

Fig. 55. Young specimen (No. 5), dorsal view.

Fig. 6. Young specimen (No. 1), lateral view.

Figs. 7 and 8. Anodontites crispata Bruguiere, from Rio de la Paila, Paila, U. S.
of Colombia. Cam. Mus. Cat. No. 61.9274 (See also Plate XLI, figs. 2 and 3).

Fig. 7a. Half-grown male (No. 8), lateral view.

Fig. 75. Half-grown male (No. 8), dorsal view.

Fig. 8. Nearly adult female (No. 10), lateral view.

Memoirs Carnegie Museum, Vol, VIII,

Plate XL

PrISODON, MoNOCONDYL.EAj Anodontites.

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656

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XLI.

Shells of Monocondylaea hollandi Ortmann, Anodontites crispata BrugiI:re,

AND Anodontites clessini (Fischer).

All figures natural size.

Fig. 1. Monocondylcea hollandi Ortmann, from Rio Guapore, near Rio Sao Simao,
Matto Grosso, Brazil. Cam. Mus. Cat. No. 61.5846.

Fig. la. Holotype, male, lateral view.

Fig. Ih. Holotype, male, dorsal view.

Fig. Ic. Holotype, male, inner view of right valve.

Figs. 2 and 3. Anodontites crispata Bruguiere, from Rio de la Paila, Paila, U. S.
of Colombia. Cam. Mus. Cat. No. 61.9274 (See also Plate XL, figs. 7 and 8).

Fig. 2a. Nearly adult specimen (sex ?), lateral view.

Fig. 25. Nearly adult specimen (sex ?), inner view of right valve.

Fig. 3. Young specimen (sex ?), lateral view.

Fig. 4. Anodontites clessini (Fischer), from Rio Vaccahy-mirim, Santa Maria,
Rio Grande do Sul, Brazil. Cam. Mus. Cat. No. 61.5820 (See also Plate XLII, figs. 1
and 2). Male (No. 12), lateral view (See also Plate XLII, fig. 1).

Memoirs Carnegie Museum, Vol, VIII.

Plate XLI.

Monocondyl^a, Anodontites,

.Jp'*

658

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XLII.

Shells of Anodontites clessini (Fischer), Anodontites hyrioides Ortmann,
Anodontites hasemani Ortmann, and Anodontites iheringi (Clessin).

All figures natural size.

Figs. 1 and 2. Anodontites clessini (Fischer), from Rio Vaccahy-mirim, Santa
Maria, Rio Grande do Sul, Brazil. Cam. Mus. Cat. No. 61.5820 (See also Plate XLI,
fig. 4).

Fig. 1. Alale (No. 12), dorsal view (Same specimen as that figured on PI. XLI,
fig. 4).

Fig. 2a. Female (No. 11), lateral view.

Fig. 26. Female (No. 11), inner view of right valve.

Figs. 3-5. Anodontites hijrioides Ortmann, from Rio Tapajos, Santarem, Para,
Brazil. Cam. Mus. Cat. No. 61.5829.

Fig. 3a. Largest specimen (No. 6), lateral view.

Fig. 36. Largest specimen (No. 6), inner view of right valve.

Fig. 4a. Half-grown specimen (No. 4), lateral view.

Fig. 46. Half-grown specimen (No. 4), dorsal view.

Fig. 5. Young specimen (No. 1), lateral view.

Figs. 6 and 7. Anodontites hasemani Ortmann, from Rio Paraguay, Santa Rita,
Matto Grosso, Brazil. Cam. Mus. Cat. No. 61.5832.

Fig. 6a. Half-grown male (No. 1), lateral view.

Fig. 66. Half-grown male (No. 1), inner view of left valve.

Fig. la. Gravid female (No. 3), lateral view.

Fig. 76. Gravid female (No. 3), dorsal view.

Fig. 8. Anodontites iheringi (Clessin), from Rio Vaccahy-mirim, Santa Maria,
Rio Grande do Sul, Brazil. Cam. Mus. Cat. No. 61.5815 (See also Plate XLIII, fig. 1,
and Plate XLIV, fig. 1).

Fig. 8a. Flalf-grown male (No. 1), lateral view.

Fig. 86. Half-grown male (No. 1), dorsal view.

Memoirs Carnegie Museum, Vol. VIII.

Plate XLII.

Anodontites.

660

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XLIII.

Shells of Anodontites iheringi (Clessin), Anodontites riograndensis (Von
Ihering), and Anodontites forbesiana (Lea).

All figures natural size.

Fig. 1. Anodontites iheringi (Clessin), from Vaccahy-mirim, Santa Maria, Rio
Grande do Sul, Brazil. Cam. AIus. Cat. No. 61.5815 (See also Plate XLII, fig. 8, and
Plate XLIV, fig. 1).

Adult female (No. 3), lateral view (See also Plate XLIV, fig. 1).

Figs. 2 and 3. Anodontites riograndensis (Von Ihering), from Rio de la Plata,
San Isidro, Argentina. Cam. Mus. Cat. No. 61.9279 (See also Plate XLIV, fig. 2).

Fig. 2a. Adult male (No. x), lateral view.

Fig. 26. Adult male (No. x), dorsal view.

Fig. 3. Adult, gravid female (No. z), lateral view (See also Plate XLIV, fig. 2).
Fig. 4. Anodontites forbesiana (Lea), from Rio Uruguay, Uruguayana, Rio Grande
do Sul, Brazil. Cam. Mus. Cat. No. 61.9280 (See also Plate XLIV, fig. 3). Half-
grown male (No. 8), lateral view.

Memoirs Carnegie Museum, Vol, VIII

Plate XLIII.

Anodontites.

662

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XLIV.

Shells of Anodontites iheringi (Clessin), Anodontites riograndensis (Von
Ihering), and Anodontites forbesiana (Lea).

All figures natural size.

Fig. 1. Anodontites iheringi (Clessin), from Rio Vaccahy-mirim, Santa Maria,
Rio Grande do Sul, Brazil. Cam. AIus. Cat. No. 61.5815 (See also Plate XLII, fig. 8,
and Plate XLIII, fig. 1). Adult female (No. 3), inner view of right valve (Same specimen
as that figured on Plate XLIII, fig. 1).

Fig. 2. Anodontites riograndensis (Von Ihering), from Rio de la Plata, San Isidro,
Argentina, Cam. Mus. Cat. No. 61.9279 (See also Plate XLIII, figs. 2 and 3). Adult,
gravid female (No. z), inner view of right valve (Same specimen as that figured on Plate
XLIII, fig. 3).

Fig. 3. Anodontites forbesiana (Lea), from Rio Uruguay, Uruguayana, Rio Grande
do Sul, Brazil. Cam. Mus. Cat. No. 61.9280 (See also plate XLIII, fig. 4).

Fig. 3a. Nearly adult female (No. 11), lateral view.

Fig. 35. Nearly adult female (No. 11), dorsal view.

Fig. 3c. Nearly adult female (No. 11), inner view of right valve.

Memoirs Carnegie Museum, Vol, VIII.

Plate XLIV.

life

Anodontites.

G64

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XLV.

Gills of Diplodon imitator Ortmann, Diplodon simillimus Ortmann, Diplodon
viCARius Ortmann, and Diplodon decipiens Ortmann.

iVll figures represent the left gills magnified to twice natural size, the inner gill to the left,

the outer gill to the right.

Fig. 1. Diplodon imitator Ortmann, from Rio Vaccahy-mirim, Santa Maria, Rio
Grande do Sul, Brazil. Cam. AIus. Cat. No. 61.9248.

Fig. la. Gills of male (No. 29) (Shell of this specimen figured on Plate XXXIV,

fig. 5).

Fig. Ih. Gills of female (No. 32).

Fig. 2. Diplodon simillimus Ortmann, from Rio Nhundiaquara, Alorretes, Parana,
Brazil. Cam. Mus. Cat. No. 61.9250.

Fig. 2a. Gills of male (No. 7).

Fig. 26. Gills of female (No. 23).

Fig. 3. Diplodon vicarius Ortmann, from creek (Rio Ribeira drainage). Aqua
(^uente, near Iporanga, Sao Paulo, Brazil. Cam. Mus. Cat. No. 61.9251. Gills of
female (No. 14).

Fig. 4. Diplodon decipiens Ortmann, from creek (tributary to Rio Iguassu),
Serrinha, Parana, Brazil. Cam. AIus. Cat. No. 61.9253.

Fig. 4a. Gills of male (No. 2).

Fig. 46. Gills of female (No. 6) (Shell of this specimen figured on Plate XXXVI,

fig. 4).

Memoirs Carnegie Museum, Vol. VIII.

Plate XLV.

Anatomy of Dii looon

666

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XLVL

Gills of Diplodon paulista (Von Ihering), Diplodon piceus (Lea), Diplodon
HILDAS Ortmann, Diplodon burroughianijs (Lea), Diplodon mogymirim
Ortmann, Diplodon berthas Ortmann, and Diplodon enno Ortmann.

All figures represent left gills magnified to twice natural size, the inner gill to the left,

the outer gill to the right.

Fig. 1. Diplodon paulista (Von Ihering), from Pio Tiete, Alogy das Cruzes, Sao
Paulo, Brazil. Cam. Mus. Cat. No. 61.9294. Gills of female (No. 4).

Fig. 2. Diplodon piceus (Lea), from Rio Uruguay, LTruguayana, Rio Grande do
Sul, Brazil. Cam. VIus. Cat. N-o. 61.5S62. Gills of female (No. 7).

Fig. 3. Diplodon hilda’ Ortmann, from Rio Jacuhy, Cachoeira, Rio Grande do
Sul, Brazil. C\arn. Mus. Cat. No. 6L5<S64. Gills of female (No. 15) (Shell of this
specimen figured on Plate XXXVII, fig. 2).

Fig. 4. Diplodon hurroughianus (Lea), from pond along Rio Negro, Santa Isabel,
Uruguay. Cam. Mus. Cat. No. 01.5859. Gills of female (No. x).

Fig. 5. Diplodon mogymirim Ortmann, from creek (tributary to Rio Alogy Guassu)
Mogy Mirim, Sao Paulo, Brazil. Cam. IMus. Cat. No. 61.9260.

Fig. 5a. Gills of male (No. 28).

Fig. 56. Gills of young female (No. 18).

Fig. 5c. Gills of adult female (No. 44).

Fig. 6. Diplodon herthce Ortmann, from Rio Jacuhy, Cachoeira, Rio Grande do
Sul, Brazil. Cam. VIus. Cat. No. 61.5805, Gills of female (No. 21).

Fig. 7. Diplodon enno Ortmann, from Rio Grande (Sao Francisco drainage),
Boqueirao, Bahia, Brazil. Cam. VIus. Cat. No. 61.9264.

Fig. 7a. Gills of young female (No. 7).

Fig. 76. Gills of nearly adult female (No. 2) (Shell of this specimen figured on

Plate XXXVIII, fig. 8).

Anatomy of Diplodon

008

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XLVII.

Left gills of Diplodon deceptus (Simpson), Castalla nehringi Von Ihering,

■ AND RIGHT GILLS OF MONOCONDYLiEA LENTIFORMIS LeA AND AnODONTITES
CLESsiNi (Fischer); twice natural size; inner gill to the left,
outer gill to the right (Figs. 1-4).

Horizontal sections of gills of Diplodon hasemani Ortmann, Diplodon imitator
Ortmann, and Diplodon decipiens Ortmann; much enlarged; inner
gill to the left, outer gill to the right (Figs. 5-7).

Fig. 1. Diplodon deceptus (Simpson), from Rio Jacuhy, Cachoeira, Rio Grande
do Sul, Brazil. Cam. Mus. Cat. 61.5868. Gills of female (No. 9).

Fig. 2. Castalina nehringi Von Ihering. from Rio Tiete, Itapura, Sao Paulo, Brazil.
Cam. AIus. Cat. No. 61.5120. Gills of half-grown female (No. 1).

Fig. 3. Monocondylcea lentiforniis Lea, from Rio Uruguay, Uruguayana, Rio
Grande do Sul, Brazil. Cam. VIus. Cat. No. 61.5841.

Fig. 3a. Gills of male (No. 1).

Fig. 36. Gills of female (No. 4).

Fig. 4. Anodontites clessini (Fischer), from Rio Vaccahy-mirim, Santa Maria,
Rio Grande do. Sul, Brazil. Cam. Mus. Cat. No. 61.5820.

Fig. 4a. Gills of male (No. 12) (Shell of this specimen figured on Plate XLI, fig. 4,

and Plate XLII, fig. 1).

Fig. 46. Gills of female (No. 11) (Shell of this specimen figured on Plate XLII,
fig- 2).

Fig. 5. Diplodon hasemani Ortmann, from Rio Guapore, near Rio Sao Simao,
Alatto Grosso, Brazil, Cam. Mus. Cat. No. 61.5857. Section of gills of gravid female
(No. 5).

Fig. 6. Diplodon imitator Ortmann, from Rio Vaccahy-mirim, Santa Maria, Rio
Grande do Sul, Brazil. Cam. Mus. Cat. No. 61.9284. Section of gills of female (No.
33) (Shell of this specimen figured on Plate XXIV, fig. 7).

Fig. 7. Diplodon decipiens Ortmann, from creek (tributary to Rio Iguassii), Ser-
rinha, Parana, Brazil. Cam. Mus. Cat. No. 61.9253. Section of gills of gravid female
(No. 5).

Memoirs Carnegie Museum, Vol, VIII.

Plate XLVII.

BMia ;

Anatomy of Diplodon, Castalia, Monocondyl®a, Anodontites.

670

MEMOIRS OF THE CARNEGIE MUSEUM.

EXPLANATION OF PLATE XLVIIL

Horizontal sections of gills of Diplodon paulista (Von Ihering), Diplodon
MOGYMiRiM Ortmann, Diplodon gratus (Lea), Diplodon deceptus
(Simpson), Castalina nehringi Von Ihering, Fossula fossiculi-
FERA (D’Orbigny), Monocondyl^a minuana D’Orbigny,

AND Anodontites patagonica rubicunda (Lea) ;
much enlarged; inner gill to the left,
outer gill to the right.

Fig. 1. Diplodon paulista (Von Ihering), from creek (tributary to Rio Mogy
Guassii), Mogy Mirim, Sao Paulo, Brazil. Cam. Mus. Cat. No. 61.9256. Section of
gills of gravid female (No. 3).

Fig. 2. Diplodon mogijmiri'm Ortmann, from creek (tributary to Rio Mogy Guassii),
Mogy Mirim, Sao Paulo, Brazil. Cam. AIus. Cat. No. 61.9260.

Fig. 2a. Section of gills of female (No. 45).

Fig. 25 Section of gills of gravid female (No. 4).

Fig. 3. Diplodon gratus (Lea), from Rio Uruguay, Uruguayana, Rio Grande do
Sul, Brazil. Cam. Mus. Cat. No. 61.5866.

Fig. 3a. Section of gills of male (No. 19).

Fig. 35. Section of gills of female (No. 20).

Fig. 4. Diplodon deceptus (Simpson), from Rio Jacuhy, Cachoeira, Rio Grande
do Sul, Brazil. Cam. Mus. Cat. No. 61.5868. Section of gills of half-grown female
(No. 6) (Shell of this specimen figured on Plate XXXIX, fig. 4).

Fig. 5. Castalina nehringi Von Ihering, from Rio Tiete, Itapura, Sao Paulo, Brazil.
Cam. VIus. Cat. No. 61.5120. Section of gills of gravid female (No. 5).

Fig. 6. Fossula fossiculifera (D’Orbigny), from Rio Tiete, Itapura, Sao Paulo,
Brazil. Cam. Mus. Cat. No. 61.5840. Section of gills of female.

Fig. 7. Monocondylcea minuana D’Orbigny, from Rio Uruguay, Uruguayana,
Rio Grande do Sul, Brazil. Cam. Mus. Cat. No. 61.5848.

Fig. 7a. Section of gills of male (No. 5).

Fig. 75. Section of gills of gravid female (No. 6).

Fig. 8. Anodontites patagonica rubicunda (Lea), from Rio Cacequy, Cacequy,
Rio Grande do Sul, Brazil. Cam. Mus. Cat. No. 61.5810. Section of gills of female.

Memoirs Carnegie Museum, Vol. VIII.

Plate XLVIII.

Anatomy of Diplodon, Castalina, Fossula, Monocondylasa, and Anodontites.

INDPJX.

aberti, Cyprogenia, 220
Abies grandis, 396
lasiocarpa, 396
Acer, 389, 397, 390, 424
macrophyllum, 423
oregonianum, 388, 423, 448
Actinonaias, 207
gibba, 237

ligamentina, 232, 234, 236, 237, 285,
294, 322

acuticosta, Castalia, 554-557
sesopus, Plethobasus, 68
sethiops, Unio, 516
affinis. Sequoia, 401, 402
alabamensis. Primus, 420
Strophitus, 206

Alasmidonta, Key to the Subgenera of, 172
Alasmidonta costata, 115, 125
heterodon, 175

marginata, 181, 184, 186, 188, 190,
191, 194

susquehannse, 181, 187, 192
varicosa, 181, 188, 190, 193, 194
alata, Proptera, 134, 252, 254-256
alatus, Unio, 252, 255
Prisodon, 562
alnifolia, Rhamnus, 396
Alnus, 389, 390, 396
carpinoides, 414
glabrata, 414

glutinosa var. denticulata, 415
Hollandiana, 387, 388, 411, 413, 414,
420

incana, 415
microdonta, 415
microdontoides, 388, 414-416
oblonga, 415
rhombifolia, 414
rugosa, 414-416

serrulata, 415
fossilis, 414
amazonia, Hyria, 562
ambigua, Castalia, 553
Simpsoniconcha, 136
Ambloma costata, 25, 33
olivaria, 229
plicata, 25-28, 53
americana, Dianthera, 264, 312
americanus, Juncus, 102, 132, 151
Amygdalonaias, Key to the Species of, 238
donaciformis, 238, 241
truncata, 238, 242, 328
angustifolia, Populus, 396
Anona, 394

Anodon reticulatus, 591
crassus, 612
trapezeum, 612

Anodonta, Key to the species and varieties
in Pennsylvania, 137
benedictensis, 144, 149
cataracta, 137, 146, 149, 152-160, 171
cygnea, 154
edentula, 197
exoticus, 624
ferussaciana, 165
fluviatilis, 160
gigantea, 145, 150
grandis, 137, 138, 142-154, 158

footiana, 137, 144, 147-152, 155,
171

gigantea, 137, 140, 145, 150, 154
harpethensis, 140
sahnonia, 140
implicata, 137, 159, 160
lacustris, 141
latoiharginata, 615
lewisi, 138, 140
lingulata, 606, 608, 609

671

672

MEMOIRS OF THE CARNEGIE MUSEUM.

marginata, 141
maryattana, 148
membranacea, 612, 613, 618
mortoniana, 606, 608
newtonensis, 160
ohiensis, 137, 162
salmonia, 140
subcylindrica, 152, 171
tryoni, 154, 155
iindulata, 195, 197
iiruguayensis, 612
virgulata, 154, 155, 159
weddelli, 606, 608
williamsi, 154, 155, 159

Aiiodonta (South American) related to
African forms of Mutela, Spatha,
453

Anodontinse, Key to the Genera of, 114;
Alasmidonta, 115; Anodonta, 115;
Anodontoides, 115; Lasmigona, 115;
Strophitus, 115

Anodontites, Key to the groups of, 588
Key to some forms of, 611
bahiensis, 622
clessini, 589, 591

crispata, 452, 588, 589, 593, 619
( Lamproscapha) ensiformis
felix, 618

forbesiana, 624, 627, 628
hasemani, 609
hohenackeri, 597
hyrioides, 604
iheringi, 611, 615, 620, 621
juparana, 597
limnoica, 619
moricandi, 623
mortoniana, 607, 608
napoensis, 588, 591
obtusa, 596, 610
pastasanus, 597
patagonica, 611, 612, 617-619
rubicunda, 611, 616, 618
puelchana, 611, 619, 620
radiata, 623

riograndensis, 624, 626, 628, 629

rioplatensis, 629
rotunda, 598, 600
spixi, 600

tenebricosa, 588, 589, 592
trapezea, 597, 598, 600
trapesialis, 621
anserina, 622
scripta, 622
trigona, 601, 605

Anodontoides, Key to the forms of, 165
buchanensis, 166

ferrusacianus buchanensis, 165, 166,
169, 170, 199
modestus, 168

approximata, Quercus, 388, 419, 420
Aralia, 398

cachemirica, 425
cordata, 425
longipetiolata, 388, 424
racemosa, 425, 426
arctica, Celastrus, 423
Populus, 408
Ardisia, 417
areolata, Quercus, 418
Aristolochia cordifolia, 407
Asia and Australia connected at one time,
455

Aspidiophyllum, 424
Aspidon, 459
australis, Diplodon, 453

bahiensis, Anodontites, 622
balsamifera, Populus, 409
balsamoides, Populus (?), 387
balzani, Iheringella, 569
baro, Castalia, 554-557
benedictensis, Anodonta, 144, 149
Betula, 389, 390, 413,
lenta, 412

multinervis, 387, 388, 411, 419, 420
papyrifera, 396

Bibliography: Naiades of Pennsylvania,
335-343

of South America, 639-641
blainyilleana, Leila, 638

INDEX.

673

borealis, Lampsilis, 292
Unio, 295

brevifolia, Sequoia, 400
brownianus, Prisodon, 564
buchanensis, Anodontoides, 166

cachemirica, Aralia, 425
cardium, Lampsilis, 301
cariosa, Lampsilis, 282, 301, 303, 313, 315,
. 318, 319, 320
cariosus, Unio, 313, 314, 316
carpinoides, Alnus, 414
Carpinus, 387, 412
Cassine, Ilex, 421
Castalia, 548

acuticosta, 554, 556, 557
ambigua, 553
baro, 554, 555, 557
hanleyana, 554-557
inflata, 558, 559
juruana, 558
pectinata, 557-559
turgida, 554
undosa, 559

Castalina martensi, 548, 551
nehringi, 463, 548
psammoica, 551
undosa, 548
Castanea, 393, 420
castelnaudi, Leila, 638
Prisodon, 564, 565
Catalpa crassifolia, 407
cataracta, Anodonta, 137, 146, 149, 152,
153, 155, 156, 157, 159, 160
catillus, Unio, 76
Celastrus, 389, 397
arctica, 423
fraxinifolius, 422
Lindgreni, 422, 423
parvifolius, 388, 422
Chamsecyparis Ehrenswardi, 405
Chara, 102
Chinchonidium, 387.
chlorophylla, Quercus, 418
cicatricosus, Plethobasus, 61-62

cincinnatiensis, Truncilla, 333
cinereoides, Quercus, 426
circulus (subrotunda), Obovaria, 226
clava, Pleurobema, 69, 86, 88, 89
clessini, Anodontites, 589, 591
Mycetopoda, 633
clintonensis, Ptychobranchus, 213
coccineus, Unio, 76
cognatus, Unio, 238
coloradensis, Heyderia, 404
communis, Juniperus, 402
complanata, Lasmigona, 133, 135,

254

complanatus, Unio, 132
Elliptio, 497

connasaugaensis, Strophitus, 206
conspicua, Lampsilis, 294
contraryensis, Quadrula, 42
cooperianus, Plethobasus, 62-65
cordata, Aralia, 425
cordifolia, Aristolochia, 407
Cornus, 387

stolonifera, 396
corrugata, Hyria, 561, 562
corymbosum, Vaccinium, 426
costata, Alasmidonta,115, 125
plicata, Amblema, 25-33
Lasmigona, 125, 132
Couttsise, Sequoia, 401, 402
crassifolia, Catalpa, 407
Juglans, 411
Credneria, 424

crispata, Anodontites, 452, 588, 593
cuneata, Populus, 408, 409
cuprea, Obliquaria, 111, 112
cupreus, Elliptio, 91, 110, 112, 272
cygnea, Anodonta, 154
cylindrica, Quadrula, 52-56
Cyperacese, 406
Cyperacites, 388, 396, 406
cyphya, Obliquaria, 61
cyphyus, Plethobasus, 65-69, 333
Cyprogenia, 217
aberti, 220
stegaria, 218

674

MEMOIRS OF THE CARNEGIE MUSEUM.

Daphnea, Quercus, 418
Decurambis marginata, 181
siisquehannse, 181, 187
varicosa, 181, 190
decurrens, Hej^deria, 404
Libocedrus, 404
delodonta, Unio, 518
deltoides, Populus, 408
Dexion, 459
Dianthera, 137

americana, 264, 312

dilatatus, Elliptio, 91, 95, 97, 100, 103, 105,
109, 263, 276
sterkii, Elliptio, 103
subgibbosus, Elliptio, 99
Eurynia, 100
Diospyros, 394, 420
Diplodon acutirostris, 505, 506
sethiops, 509, 530
ampullaceus, 518
aplatus, 487
apprimus, 520, 512, 513
australis, 453
berthse, 528

burroughianus, 483, 517, 518
charriianus, 504, 506, 509, 515, 517
chilensis, 487, 504
coriaceus, 485 '
deceptus, 539, 540
decipiens, 494, 499
ellipticiim, 471
ellipticus, 526-532
santanus, 527
enno, 531
expansus, 485
faba, 509
felipponei, 520
firmus, 511, 512
fontainianus, 537-539, 542
fortis, 506, 509
frenzeli, 488, 489
funebralis, 545
gassiesi, 487
granosiis, 484, 485, 497
granuliferus, 486

gratus, 534, 539, 540-542
deceptus, 534, 539, 540
greeffea'nus, 523
guaranianus, 474, 476, 479, 483
hasemani, 478, 479, 483
hidalgoi, 506, 509
hildse, 514, 516
huapensis, 488-490
hylseus, 473-477
imitator, 491, 495-500
lacteolus, 513, 517, 518
lepidus, 511
martensi, 491, 497
mimus, 486, 497
modestus, 497
multistriatus, 485
mogymirim, 520, 523
nocturnus, 545
parallelopipedon, 483, 504
paranensis, (Cyclomya) 526, 542, 546
parcus, 506, 509
patagonicus, 488
paulista, 501, 503
piceus, 510, 511
piger, 518
piracicabana, 507
psammactinus, 485
rotundus, 531, 541
santamarise, 495
semigranosus, 486
simillimus, 494-502
solidulus, 487
suavidicus, 524
suppositus, 503
trifidus, 473, 479, 480, 483
Uruguay ensis, 512, 513
trivialis, 520, 521
vicarius, 494, 497, 501, 502
wagnerianus, 527
wymani, 512, 513, 518-520
Diplodon, Final remarks on the genus,
547-548

Key to groups in, 472-473
subdivisions, 471
dolobrata, Thuyopsis, 404

INDEX.

675

donaciformis, Amygdalonaias, 238, 241
dorfeuillana, Quadrula, 36-38
Douglass, Earl: collection of fossil plants,
385

dubium, Taxodium, 403

Ecological conditions indicated by the
fossil flora from Missoula, Mon-
tana, 395

Elliptio, Key to the Pennsylvanian species
and varieties, 91
complanatus, 497
cupreus, 91, 110, 112, 272
dilatatus, 91, 95, 97, 100, 103, 105,
109, 263, 278
sterkii, 91, 98, 101, 103
subgibbosus, 99
fisherianus, 91, 112, 113, 272
niger, 91, 94

violaceus, 91, 103, 106, 108-112, 276
Ebenaceae, 420
Echinopanax horridum, 396
edentula, Anodonta, 197
edentulus, Strophitus, 167, 196, 197, 199,
204-206

Ehrenswardi, Chamsecyparis, 405
elaena, Quercus, 418
elaeomorpha, Quercus, 418
Ellipsaria, 207

fasciolaris, 97, 208, 213
occidentalis, 213
elliptica, Quercus, 418
ellipticum, Diplodon, 471
Engelmanni, Picea, 396
ensiformis, Anodontites, 630
Epaspidon, 459
Epidexion, 459
Equisetites, 398, 399
Equisetum., 398
hyemale, 399
insculptum, 388, 398, 399
variegatum var. Jesupi, 399
wyorningense, 399
Ericaceae, 422

eriganensis, costata, 131 (Lasmigona)

esula, Leila, 638

europaeus, Glyptostrobus, 387, 401
Eurynia, Key to the species of the sub-
genus, 270

fabalis, 258, 259, 261, 263, 266, 267
iris, 234, 263, 266, 267, 269, 270,
285

nasuta, 270, 271
exasperata, Hyria, 562
Expedition of Carnegie Museum to Cen-
tral South America, 451
Explanation of plates representing fossil
plants from beds of volcanic
ashes near Missoula, Mon-
tana, 428-450

Ditto, Naiades of Pennsylvania,
344-384

Ditto, Naiades of South America,

642-670

\

fabalis, Eurynia, 258, 259, 261, 263, 266,
267

Unio, 261

fasciolaris, Ellipsaria, 97, 208, 213
Obliquaria, 207

fasciola, Lampsilis, 282, 303, 309, 312
fatuus, Lampsilis, 267
felix, Anodontites, 618
ferussaciana, Anodonta, 165
Anodontoides, 165, 199
buchanensis, Anodontoides, 166, 169,
170

modestus, Anodontoides, 168
Ficus, 389, 394, 396

(?) prunifolia, 388, 420
fisherianus, Elliptio, 91, 112, 113, 272
Unio, 110

flava, Fusconaia, 7, 14-19
flexuosa, Quercus, 388, 418
fluviatilis, Anodonta, 160
forbesiana, Anodontites, 624-628, 629
fossiculifera, Fossula, 571 z
Fossil flora from Winston, Montana, Re-
marks on, 398
Fossula fossiculifera, 571

676

MEMOIRS OF THE CARNEGIE MUSEUM.

Fossil Flora from Missoula, Montana,
Ecological conditions in-
dicated by, 395-398

Relations to the nearest Eo-
cene and Miocene Floras
of the West, 387-395
fragilis, Paraptera, 247, 253, 256
Unio, 247
fulgidiis, Unio, 76
fuliginosus, Unio, 106
furcinervis americana, Quercus, 419, 421
Ilex, 388, 421, 422
Quercus, 419
Fusconaia, 71

Key to the species and varieties, 7
flava, 7, 14-19

parvula, 7, 21-25
trigona, 7, 19-21
subrotunda, 7-11, 70

kirtlandiana, 7, 11-14

genetrix, Populus, 408
gesneri, Strophitus, 206
gigantea, Anodonta, 145, 150
Glabaris, 453

simpsonianus, 626
trapesialis cygneiformis, 626
wymani, 615
glabrata, Alnus, 414
glana, Unio, 257
glandulifera, Populus, 408
glutinosa var. denticulata, Alnus, 415
Glyptostrobus europseus, 387, 401
gracilis, Thuyopsis, 388, 392, 403, 404
grandis, Abies, 396
Grewiopsis tennesseensis, 407
grandis, Anodonta, 137, 138, 142, 145, 146,
148-151, 153, 154, 158
footiana, Anodonta, 137, 144, 147-
152, 157, 171

gigantea, Anodonta, 137, 140, 154
salmonia, Anodonta, 140
greeffeanus, Unio, 527
guarayana, Monocondylsea, 572
gubernaculum, Truncilla, 334

Unio, 331

hanleyana, Castalia, 554-557
harpethensis, Anodonta, 140
hasemani, Anodontites, 609
Haydenii, Hypnum, 400-402
Juniperus (?), 402
Sequoia, 387-400, 402, 411, 419
Hedera, 393
Heteranthera, 264
heterodon, Unio, 172
heterophylla, Tsuga, 396
Heyderia coloradensis, 404
decurrens, 404
Hicoria, 414

higginsi, Lampsilis, 323, 324
hohenackeri, Anodontites, 597
Hollandiana, Alnus, 387, 388, 411, 413,
414, 420

hollandi, Monocondylsea, 585
horridum, Echinopanax, 396
hyalinus, Unio, 125
hydiana, Lampsilis, 287, 289
hyemale, Equisetum, 399
Hypnum Haydenii, 400-402
Hyria amazonia, 562
corrugata, 561, 562
exasperata, 562
rugosissima, 562
transversa, 562, 563
Hyridella, 453

Hyriinse, genera of : Diplodon, Castalina,
Castalia, Castaliella, Callonaia,
Hyria, Prisodon, 460-468
Hyriinse 457-460
Hyriine glochidia, 468-471
hyrioides, Anodontites, 604

Icacorea, 417
Iheringella balzani, 569
iheringi, Anodontites, 611, 615
Ilex, 397

Cassine, 421

furcinervis, 388, 421, 422
longifolia, 421

INDEX.

677

imbricaria, Quercus, 421
implicata, Anodonta, 137, 159, 160
infiana, Alnus, 415
incrassatus, Unio, 94
inermis, Monocopdylsea, 585
inflata, Castalia, 558, 559
insculptum, Equisetum, 388, 398, 399
iris, Eurynia, 234, 263, 266, 267, 269, 270,
285

jejunus, Unio complanatus, 105
Jennings, Grace K. : photographic repro-
ductions, 386

Jennings, 0. E., Fossil Plants from the
Beds of Volcanic Ash near Miss-
oula, Western Montana, 385-450
Juglans, 389, 397, 414
crassifolia, 411
pentagona, 388, 410, 411
rugosa, 411

Juncus americanus, 102, 132, 151, 270
Juniperus communis, 402
(?) Haydenii, 402
scopulorum, 397
virginiana, 397
juparana, Anodontites, 597
juruana, Castalia, 558

kieneriana, Quadrula, 37
kirtlandiana, Obovaria, 221
‘‘kleineriana,” 37

lacustris, Anodonta, 141
Lampsilinse, Key to the genera of, 206:
Actinonaias, 207 ; Amygdalonaias,
207 ; Cyprogenia, 207 ; Ellipsaria,
206; Eurynia, 207; Lampsilis, 207;
Obliquaria, 207 ; Obovaria, 207 ; Par-
aptera, 207 ; Plagiola, 207 ; Proptera,
207 ; Toxolasma, 207 ; Truncilla,
207

Lampsilis, Key to the species and varie-
ties of, 282
borealis, 292
cardium, 301

cariosa, 282, 301, 303, 313, 314, 315,
316, 318-320
conspicua, 294
fasciola, 282, 303, 309, 312
fatuus, 267
higginsi, 323, 324
hydiana, 287, 289
ligamentina, 322

luteola, 234, 266, 267, 282, 283, 285,
288, 289, 290, 292-294, 297
rosacea, 282, 289, 290
occidentalis, 213
ochracea, 282, 303, 318
ochraceus, 296, 316
orbiculata, 303, 320, 322, 323, 324,
282

ovata, 282, 297, 298, 303, 301, 306
canadensis, 282, 306, 307, 309
lurida, 308

ventricosa, 282, 298, 299, 301,
303, 306, 307, 309, 312, 313,
315

radiata, 282, 292, 294, 318
rosacea, 286, 291, 292
superiorensis, 291, 292
Lamproscapha, 629, 630
lanceolatus, Unio, 112
Langsdorfii, Sequoia, 387, 400, 403
Larix occidentalis, 396
lasiocarpa, Abies, 396
latomarginata, Ancdonta, 615
Lasmigona, Key to the subgenera of, 115
Key to the forms of, 125
complanata, 135, 254
costata, 125, 132
eriganensis, 131
(Platynaias), 115

subviridis, 115, 117, 121
viridis, 115, 116
plebeia, 120
(Pterosyna), 115
complanata, 133
viridis, 121, 124, 136
Lasmonos, 247
latissima, Eurynia, 279

678

MEMOIRS OF THE CARNEGIE MUSEUM.

Lauraceie, 417, 420
laiirifolia, Querciis, 417
laiiriformis, Qiiercus, 418
laiirina, Quercus, 417
laiiroide.s, Quercus, 417
laurophylla, Quercus, 417
laurosimulans, Quercus, 388, 416
I.aurus, 393

princeps, 417

legumen, Mycetopoda, 633
Leguminosa3, 422
Leila, 458, 587, 637
castelmani, 638
esula, 638
blainvilleana, 638
pulvinata, 638
spixi, 638

lens (levigata), Obovaria, 226
lenta, Betula, 412

leiitiformis, Monocondylsea, 573, 578, 581,
587

leptodon, Lbiio, 247
Lesquereuxii, Typha, 388, 389, 405
Leucsena, 394
levigata, Obovaria, 220

Obovaria subrotunda, 221
lewisi, Anodonta, 138, 140
Libocedrus decurrens, 404
liganientina, Actinonaias, 232, 234, 236,
237, 285, 294, 322
gibba, Actinonaias, 237
Lampsilis, 322

liinnoica, Anodontites, 618, 619
Lindgreni, Celastrus, 423
lineolata, Obliquaria, 243
Plagiola, 243

lingusefolia, Sabina, 388, 389, 404, 405
lingulata, Anodonta, 606-609
lividus, Unio, 257
longifolia. Ilex, 421
longipetiolata, Aralia, 388, 424
Lyelli, Quercus, 418

luteola, Lampsilis, 234, 236, 267, 282, 283,
285, 288-297

rosacea, Lampsilis, 282, 289

macrophyllum, Acer, 423
Margaritana margaritifera, 2-6, 154
marginata, 190
Margaritanidae, 2

margaritifera, Margaritana, 2-6, 154
Mya, 2

marginata, Alasmidonta, 181, 184, 186,
188, 190-194
Anodonta, 141
Margaritana, 190
susquehannse, Alasmidonta, 192
martensi, Castalina, 548, 551
Unio, 491

maryattana, Anodonta, 148
membranacea, Anodonta, 012, 613, 618
metaneVra, Obliquaria, 33
Quadrula, 47-52
wardi, Quadrula, 48, 50
microdonta, Alnus, 415
microdontoides, Alnus, 388, 414, 415, 416
Micromya, Key to the forms of, 261
fabalis, 261, 269
iris, 261, 265

iris novi-eboraci, 26, 267, 268
nasuta, 273
nebulosa, 269

recta, 97, 270, 276, 280, 281
sageri, 279
sageri latissima, 279
subrostrata, 276

minuana, Alonocondylsea, 579, 583

parchappi, Monocondylsea, 580-583
Monocondylsea, Key to the species of, 573
guarayana, 572
hollandi, 585
inermis, 585

lentiforniis, 573, 578, 581, 587
minuana, 579, 583

parchappi, 580-583
obesa, 583

paraguayana, 577, 581
pazi, 581

Monodonta undulata, 172, 176
monticola, Pinus, 396
moricandi, Anodontites, 623

INDEX.

679

mortoniana, Anodontites, 607, 608
mucronata, Pseudotsuga, 396
multinervis, Betula, 387, 388, 411, 419, 420
Murrayana, Pinus, 396
Mussels (freshwater) of South America,
Naiades of South America. By A.
E. Ortmann, 451-670
Mutelidse, 453, 455
Mutelinse, 453, 457
African, 637

Found in South America east of the
Andes, Pacific drainage in Ecuador,
Central America to Southern Mexi-
co, 454

General remarks on, 565-567
Genera of the South American, 567
Key to genera of South American, 568
Species of South American, 569
South American, 637
Mya margaritifera, 2
Mycetopoda, 587, 632
clessini, 633
legumen, 633
siliquosa, 633, 634
subsinuata, 635
staudingeri, 635
Myrica, 390
Myrsinacese, 417
Myrsihe, 417
Mytilus cygneus, 137

Naiades; Affinities of South American
forms, 451

Bibliography of, 335-343, 639-641
Explanation of plates, 344-384, 642-
670

Collected by Haseman, Eigenmann,
and Windhausen, 452
Contribution to knowledge of South
American. By A. E. Ortmann,
451-670

Diagnostic characters of South Amer-
ican, 456

Difficulties encountered in recogniz-
ing described forms, 452

General remarks on affinities and
geographical distribution of S. Am-
erican, 453-455

Important specific characters not
heretofore emphasized, 452
Largest collection of South American
Naiades ever secured, 451, 452
Monograph of the Naiades of Penn-
sylvania. Part III. Systematic
account of the genera and species.
By A. E. Ortmann, 1-384
Spelling of the word, 1
South American placed by early
writers in Unio, Anodonta, Mono-
condylsea, Hyria, Castalia, 453
Systematic arrangement and geo-
graphical distribution of S. Ameri-
can, 454

Two groups in South America, 454
napoensis, Anodontites, 588
nasuta, Eurynia, 273, 112
nebulosa, Eurynia, 269
Nectandra, 417
nehringi, Castalina, 463, 548
neriifolia, Quercus, 418
newtonensis, Anodonta, 160
niger, Elliptio, 91, 94
nigra, Unio, 91, 93
nocturnus, Unio, 545
northamptonensis, Unio, 105, 106

obesa, Monocondylaea, 583
Obliquaria cuprea. 111, 112
cyphya, 61
fasciolaris, 207
lineolata, 243
metanevra, 33
reflexa, 213, 214
tuberculata, 56

obliquum catillus, Pleurobema, 69, 74, 75,
79, 81, 82, 85

coccineum, Pleurobema, 80
pauperculum, Pleurobema, 80
Pleurobema, 69, 70, 73, 74, 77, 81,
84-86

680

MEMOIRS OF THE CARNEGIE MUSEUM.

obliqiius, Prisodon, 564, 565
oblonga, Alnus, 415
oblongifolia, Sequoia, 388, 400
Obovaria, Key to the subgenera of, 220
circulus (subrotunda), 226
lens (levigata), 223, 226
(Pseudoon) olivaria, 229
(Unio) retusa, 220
retusa, 221
stegaria, 217

subrotunda, 223, 225, 227
levigata, 221
obtusa, Anodontites, 596
occidentalis, Ellipsaria, 213
Lampsilis, 213
Larix, 396
Unio, 213
Taxodium, 387

ochracea, Lampsilis, 282, 296, 303, 316,
318

ochraceus, Unio, 292
ohiensis, Anodonta, 137, 162
Olafseni, Quercus, 419
Oligocene flora of Missoula, Montana,
Descriptions of species, 398-427
Winston, Montana, Descriptions
of species, 398-427
olivaria. Ambloma, 229
Olriki, Taxites, 406
oratus, Unio, 318

orbiculata, Lampsilis, 282, 303, 320-324
oregonianum, Acer, 388, 423
Ortmann, Arnold E., A Monograph of the
Naiades of Pennsylvania. Part III.
Systematic Account of the Genera
and Species, 1-384.

South American Naiades; A Contri-
bution to the Knowledge of the
Freshwater Mussels of South
America, 451-670
Ostrya virginiana, 412
ovata canadensis, Lampsilis, 282, 306-309
Lampsilis, 282, 297, 298, 301, 303,
306

lurida, Lampsilis, 308

ventricosa, Lampsilis, 282, 298, 299,
301, 306, 307, 309, 312, 315

palaeocorymbosum, Vaccinium, 388, 426
paleomelas, Populus, 408
papyrifera, Betula, 396
paraguayana, Monocondylsea, 577, 581
paraguayensis, Unio, 546
Paraptera fragilis, 247, 253, 256
parva, Fusconaia, 7, 21-25
parvifolius, Celastrus, 388, 422
parvum, Toxolasma, 258, 263
pastasanus, Anodontites, 597
X patagonica, Anodontites, 611, 612, 617,
618

rubicunda, Anodontites, 611, 616, 618
patagonicus, Unio, 488
pauperculum, Pleurobema obliquum, 69
83, 86

pauperculus, LTnio, 83
pazi, Anodontites, 618
Monocondylsea, 581
pectinata, Castalia, 557-559
peculiaris, Unio, 510
pentagona, Juglans, 388, 410, 411
percoarctatus, Unio, 108
perplexa, Truncilla, 332, 334
Persea, 394
pertenuis, Unio, 125
Picea Engelmanni, 396
Pinus monticola, 396
Murrayana, 396
ponderosa, 396, 397
Plagiola, 207

lineolata, 243

Plants (Fossil) from Beds of Volcanic Ash
near Missoula, Western Montana.
By 0. E. Jennings, 385-450
platania, Quercus, 419
Platynaias, Key to the Species of the Sub-
genus, 115

Plethobasus, Key to the species of, 61
aesopuS, 68
cicatricosus, 61, 62
cooperianus, 63-65

INDEX.

681

cyphyus, 65-69, 333
Pleurobema clava, 69, 86, 88, 89

catillus, 69, 74, 75, 79, 81, 82, 85
coccineum, 69, 78, 79, 80-83
cordatum, 69, 72, 74, 75
obliquum, 69, 73, 74, 77, 81, 84,
85, 86

pauperculum, 69, 80, 83, 86
riibrum, 69, 78, 79, 84, 85-
plicata, Amblema, 53
Thuya, 396
Podocarpus, 400, 406
Polytrichum, 402
ponderosa, Pinus, 396, 397
Populus, 389, 390, 407
angustifolia, 396
arctica, 408
balsamifera, 409
balsamoides (?), 387
cuneata, 408, 409
deltoides, 408
genetrix, 408
glandulifera, 408
paleomelas, 408
primseva, 408
smilacifolia, 388, 409, 410
tremuloides, 396

Zaddachi, 387, 388, 392, 406, 408,
410, 411

Potamogeton, 264
primseva, Populus, 408
Prisodon alatus, 563
brownianus, 564
castelnaudi, 564, 565
obliquus, 564, 565

Prolasmidonta heterodon, ( Alasmidonta) ,
173

Proptera alata, 134, 252, 254, 256
Protophyllum, 424
prunifolia. Ficus (?), 388, 420
Prunus alabamensis, 420
psammoica, Castalina, 551
(Pseudoon) olivaria, Obovaria, 229
Pseudotsuga mucronata, 396
Pterosyna, 132

complanata (Lasmigona), 133
Ptychobranchus clintonensis, 213
puelchana, Anodontites, 611, 619
pulvinata, Leila, 638
purpureus, Unio, 110
pustulatus, Unio (Quadrula), 39
pustulosa, Quadrula, 34-40
dorfeuillana, 3, 32
kieneriana, Quadrula, 37
schoolcraftensis, Quadrula, 38-40
pustulosus, Unio (Quadrula), 39

Quadrula, Key to the species and varie-
ties, 33

coccinea paupercula, 83
contraryensis, 42
cylindrica, 52-56
dorfeuillana, 36, 37, 38
kieneriana, 37
metanevra wardi, 47-52
pernodosa, 37
pustulatus, 39
pustulosa, 33-40
pustulosa schoolcraftensis, 38-40
quadrula, 40-43, 68
rubiginosa, 39
verrucosa, 43-47
Quercus, 389, 390, 393, 397
approximata, 388, 419, 420
areolata, 418
chlorophylla, 418
cinereoides, 426
Daphnea, 418
elsena, 418
elseomorpha, 418
elliptica, 418
flexuosa, 388, 418
furcinervis, 419

americana, 419, 421
imbricaria, 421
laurifolia, 417
lauriformis, 418
laurosimulans, 388, 416
Lyelli, 418
neriifolia, 418

682

MEMOIRS OF THE CARNEGIE MUSEUM.

Olafseni, 419
platania, 419
simiilata, 418
socia, 418

Steenstrupiana, 419

racemosa, Aralia, 425, 426
radiata, Anodontites, 623
Lampsilis, 282, 292, 294
radiatus, Unio, 293, 294
rangiana, Truncilla, 325, 331-335
recta, Eurynia, 97, 261, 270, 278-281
reflexa, Obliquaria, 213, 214
Relations of the fossil flora of Missoula to
the nearest Eocene and Miocene
floras of the west, 387
reticulatiis, Anodon, 591
retusa (Unio), Obovaria, 220
Rhamnacese, 422
Rhamniis alnifolia, 396
rhombifolia, Alnus, 414
riograndensis, Anodontites, 624-628, 629
rioplatensis, Anodontites, 629
roanokensis, Unio, 105
rosacea, Lampsilis, 292
Rosaceie, 422
rosaceus, Unio, 318
rotunda, Anodontites, 598, 600
Rotundaria tuberculata, 57-61
rubicunda, Anodontites, 616, 618
rubiginosa, Quadrula, 39
rubrum, Pleurobema obliquum, 69, 78,
79, 84, 85

rugosa, Alnus, 414-416
Juglans, 411
rugosissima, Hyria, 562
rugosum, Triplodon, 562

Sabina, 397

linguscfolia, 388, 389, 404, 405
scopulorum, 405
sageri, Eurynia, 279
Salix, 393, 396
salmonia, Anodonta, 140
Sapindus, 394

sapotalensis, LTnio, 232
scopulorum, Juniperus, 397
Sabina, 405

sempervirens. Sequoia, 402
Sequoia, 390, 397, 405
affinis, 401, 402
brevifolia, 400
Couttsiae, 401, 402

Haydenii, 387-389, 400, 402, 411, 419
Langsdorfii, 387, 400, 403
oblongifolia, 388, 400
sempervirens, 402
serrulata, Alnus, 415
fossilis, Alnus, 414
siliquosa, Alycetopoda, 633, 634
simpsonianus, Glabaris, 626
Simpsoniconcha, 136
ambigua, 136
simulata, Quercus, 418
smilacifolia, Populus, 388, 409, 410
socia, Quercus, 418
solidus, Unio, 76

South America and Australia connected
at one time, 454

South American Continent, Origin of, 454
spillmani, Strophitus, 206
spixi, Anodontites, 600
Leila, 638

Steenstrupiana, Quercus, 419
stegaria, Cyprogenia, 218
Obovaria, 217

sterkii, Elliptio dilatatus, 91, 101
stolonifera, Cornus, 396
Strophitus alabamensis, 208
connasaugaensis, 206
edentidus, 167, 196-199, 204-206
gesneri, 206
spillmani, 206
undulatus, 190, 195, 197
Styganodon, 587, 588
subcylindrica, Anodonta, 152
subrostrata, Eurynia, 276
subrotunda, Fusconaia, 7-11, 70
Obovaria, 225, 227, 228
subviridis, Lasmigona, 117, 120

INDEX.

683

Lasmigona (Platynaias), 115, 121
siiperiorensis, Lampsilis, 291, 292

Taxites, 400
Olriki, 406
Taxodium, 400
dubium, 403
occidentalis, 387
Taxus, 400

tenebricosa, Anodontites, 588, 592
tennesseensis, Grewiopsis, 407
.Terminalia, 420
“Tetragenae,” 71
Tetraplodon, 453
texasense, Toxolasma, 260
Thuya, 397

plicata, 396
Thuyopsis, 397
dolobrata, 404
gracilis, 388, 392, 403, 404
torulosa, Truncilla, 331, 334
Toxolasma parvum, 257, 258, 263
texasense, 260
transversa, Hyria, 562, 563
trapesialis, Anodontites, 621
cygneiformis, Glabaris, 626
scripta, Anodontites, 621, 622
trapezea, Anodontites, 597-600
trapezeum, Anodon, 612
tremuloides, Populus, 396
trigona, Anodontites, 601, 605
Fusconaia, 7, 19-21
Triplodon rugosum, 562
triquetra, Truncilla, 239, 325, 329, 330
truncata, Actinonaias, 239

Amygdalonaias, 238, 242, 328
Truncilla, 238
Unio, 238

Truncilla, Key to the Pennsylvanian spe-
cies of, 325
cincinnatiensis, 333
gubernaculum, 334
perplexa, 332, 334
rangiana, 325-335
torulosa, 331, 333, 334

triquetra, 239, 325, 328, 330
truncata, 238

tryoni, Anodonta, 154, 155
Tsuga heterophylla, 396
tuberculata, Rotundaria, 57-61
turgida, Castalia, 554
Typha, 396, 406

Lesquereuxii, 388, 389, 405

Ulmus, 412
undosa, Castalia, 559
Castalina, 548
undulata, Alasmidonta, 177
Anodonta, 195
Monodonta, 172, 176
undulatus, Strophitus, 190, 195, 197
Unio aethiops piracicabana, 514
alata, 252
alatus, 255
borealis, 295
catillus, 76
coccineus, 76
cognatus, 238
complanatus, 132
jejunus, 105
delodonta, 518
fabalis, 261
fasciata cuprea, 112
fisherianus, 110
fragilis, 247
fulgidus, 76
fuliginosus, 106
giana, 257
greeffeanus, 520
gubernaculum, 331
heterodon, 172
hyalinus, 125
incrassatus, 94
jejunus percoarctatus, 105
lanceolatus, 112
leptodon, 247
lividus, 257
martensi, 491
nasuta, 112
nigra, 91, 93

684

MEMOIRS OF THE CARNEGIE MUSEUM.

nocturnus, 546
northamptonensis, 105, 106
occidentalis, 213
ochraceiis, 292
oratus, 318
paraguayensis, 546
patagonicus, 488
peculiaris, 510
pertensis, 125
piirpiireus, 110
radiatus, 293, 294
recta, 261, 270
retusa, 220
roanokensis, 105
rosaceiis, 318
sapotalensis, 232
solidus, 76
truncata, 238
viridis, 115
Unionidse, 455

iiiTiguayensis, Anodonta, 612, 613
Unionidse, 6

Unioninse, Key to the genera, 6
Vacciniuin, 396, 427

corymbosum, 426
palseocorymbosum, 388, 426
Vallisneria, 264

varicosa, Alasmidonta, 188, 189, 193, 194
variegatum var. Jesupi, Equisetiim, 399
ventricosa, Larapsilis, 303
verrucosa, Qtiadrula, 43-47
Viburnum, 387

violaceus, Elliptio, 91, 103-112, 276
virginiana, Juniperus, 397
Ostrya, 412
Virgula, 587, 588

virgulata, Anodonta, 154, 155, 159
viridis, Lasmigona, 120, 121, 124, 136
Lasmigona (Platynaias), 115, 116
Unio, 115

Von Ihering, H. 452, 453

weddelli, Anodonta, 606, 608
williamsi, Anodonta, 154, 155, 159
wymani, Glabaris, 615
wyomingense, Equisetum, 399

Zaddachi, Populus, 387, 388, 392, 406,
408, 410, 411

44. Additions to Flora of Allegheny County, Pa.

Jennings 15

45. New Species of Kneifila. Jennings 05

46. Occurrence of Triglochin palustris in Pa. Jen-

nings .06

47. New Species of Ihidium. Jennings 10

48. Agate Spring Fossil Quarry. Peterson 10

49. Two New^ Birds from British E. Africa. Obeb-

HOLSER .06

50. The Chazy Formation and Its Fauna. Eay-

MONh 1.60

51. A New American Oyhele. Narraway and Eay-

MOND .15

52. Plastron of the Protosteginae. Wieland 15

53. New Species of Testudo from the Miocene, etc.

0. P, Hay 26

54. Miocene Beds of W. Nebraska and E. Wyoming

and Their Vertebrate Faunae. Peterson 1.00

65. New Species of Lonicera from Pa. Jennings. .06

56. Merycochcerus, etc. Douglass.

57. New Merycoidodonts. Douglass. (Nos. 56

and 57 sold together.) 1.00

58. Further Collections of Fishes from Paraguay.

Eigenmann, McAtee, and Ward 1.26

59. Undetermined Element in the Osteology of the

Mosasauridae. Holland 20

60. Gasteropoda of the Chazy. Eaymond 1.36

61. Occurrence of Wynea Americana in Pa. Jen-

nings 05

62. Account of Pleistocene Fauna Discovered at

Frankstown, Pa. Holland 10

63. Vertebrate Fossils from the Vicinity of Pitts-

burgh, Pa. Case 16

64. IllaenidaB from the Black Eiver Limestone near

Ottawa, Canada, Eaymond and Narraway. . .20

65. Rhinoceroses from the Oligocene and Miocene

of North Dakota and Montana. Douglass . . .26

66. Fossil Horses from North Dakota. Douglass, .30

67. Oligocene Lizards. Douglass 20

68. The Type of Stenomylus gracilis. Peterson . . .26

69. New Species of Birds from Costa Rica, etc.

Carrikeb 10

70. Costa Rican Formicariidae. Cabriker 05

71. Vertebrate Fossils from the Fort Union Beds.

Douglass 46

72. List of the Lepidoptera of Western Pa., etc.

Engel 1.26

73. Fauna of Upper Devonian in Montana. Fossils

of Red Shales. Raymond 60

74. New Species of Procamelus from Upper Mio-

cene of Montana. Douglass 30

75. Sections of the Conemaugh Series -between

Pittsburgh and Latrobe, Pa. Raymond 36

76. List of the Unionidae of Western Pa. Ortmann .50

77. Geological Reconnaissance in North Dakota,

Montana, and Idaho. Douglass 1.00

78. Botanical Survey of Presque Isle. Jennings. . 2.75

79. Sesqui-Centennial (Pittsburgh) Relics. Stewart .45

80. Dromomers^, New Genus of American Rumi-

nants. Douglass .30

81. Fossils from Glacial Drift and Devonian and

Mississippian near Meadville, Pa. Millard,. .10

82. New Species of Helodus. Eastman .05

83. Charles Chauncey Mellor. Holland 15

84. Reports of Expedition to British Guiana, etc.

Eigenmann 60

85. Reports of Expedition to British Guiana, etc.

Durbin 25

86. Odonata of the Neotropical Region, Exclusive

of Mexico and Central America. Calvert. . . 2.26

87. Deinosuchus hatcheri, a New Genus and Species

of Crocodile. Holland 20

88. Reports on Expedition to British Guiana, etc.

o Blossbb 16

89. Description of Some New Titanotheres from

the Uinta. Douglass 26

90. Annotated List of the Birds of Costa Rica In-

cluding Cocos Island. Carrikeb 3.00

91. Geology of the Coast of the State of Alagdas,

Brazil. Brannee 40

92. Fossil Fishes from the Bituminous Shales at Ri-

acho Doce, Alagoas, Brazil. Jordan 56

93. Ordovician Trilobites, No. n. Asaphidse from

the Beekmantown, Raymond 36

94. Ordovician TrEobltes, No. HI. Raymond &

Narraway 36

95. Ordovician Trilobites, No. IV. Raymond 40

96. Fishes from Irkutsk, Siberia. Jordan and

Thompson .30

97. - South American Tetrigidas. Bruner 1.00

98. Fauna of the Allegheny and Conemaugh Series

in Western Pa. Raymond 30

99. Ichthyological Survey About the San Juan

Islands, Washington. Starks 76

100. New Species of Pygidium. Eigenmann 10

101. The Brachiopoda and Ostracoda of the Chazy.

Raymond 60

102. A New Camel from the Miocene of Western

Nebraska. Peterson 16

103. Skeleton of Stenomylus hitchcocki, etc. Peter-

son .16

104. Skeleton of Diceratherium cooki. Peterson. . .16

105. Carnegie Museum Expedition to Central South

America, 1907-1910, Holland 16

106. Report Upon the Expedition of the Carnegie

Museum to Central South America. Hase-
MAN & Eigenmann 25

107. New Species of Fishes, etc., from Central South

America. Haseman 50

108. Annotated Catalog of Cichlid Fishes from Cen-

tral South America. Haseman 1.26

109. New Species of Fishes from the Rio Iguassu,

Haseman 66

110. Ornithology of the Bahama Islands. Todd &

Worthington 76

112. South American Acridoidea. I. Bruner 1.00

113. Species of Hasemania, Hyphessobrycon, and

Hemigrammus. Ellis 26

114. New Characins in the Carnegie Museum. Eigen-

mann 30

115. Jurassic Saurian Remains Ingested within Fish.

Eastman 20

116. Autograph Letter of U. S. Grant to Edwin M.

Stanton. Holland 16

117. Albert J. Barr. By W. J. Holland 10

118. Seventeen New Neotropical Birds. Todd 26

119. Dr. David Alter, the First Discoverer of Spec-

trum Analysis. Holland 10

120. Two Mummy Labels. Allen 10

121. The Families and Genera of Najades. Obt-

MANN 1.00

122. Group of Stenomylins in the Carnegie Museum.

Peterson 26

123. Tertiary Fish-remains from Spanish Q-uinea.

Eastman .26

124. Plated Nematognaths. Ellis 66

125. New Species of Camharus from the Isle of

Pines. Ortmann 15

126. Sedum Camegiei, a New Species of the Family

Crassulacese, etc. Hamet .10

127. Two New Species of Fishes from Peru. Eigen-

MANN 16

128. South American Locusts ( Acridoidea) . II.

Bruner 75

129. Revision of the Genus Chsemepelia. Todd 85

130. New Genus and Species of Abyssinian Rodents.

Childs Erick .20

131. New Titanothere from the Uinta. Peterson. . .20

132. Small Titanothere from the Lower Uinta. Peter-

son 10

133. Osteology of Lasiopyga and Callithrix, etc.

Shueeldt 25

134. New Rhynchocephalian from Solenhofen. Grier .16

135. Scales. of South American Characinid Fishes.

Cockerell 26

136. Skeleton of Platigonus Leptorhinus. Peter-

son 10

137. Lichens Collected in the Thunder Bay District,

Ontario. E. Heber Howe, Jr 10

138. Preliminary List of Fossil Plants in the Roof

of the Pittsburgh Coal. Grier 10

139. Undescribed Remains of the Uinta Titanothere

Dolichorhinus. Peterson ,16

140. Triassic Fishes Belonging to the Catopteridse

and Semionotidse. Eastman .16

141. Osteology of Promerycochoerus. Peterson ... .40

142. Correction of a Generic Name. Peterson 05

143. Skull of Bison Crassicornis. Holland .10

144. Serrasalminse and Mylinse. Eigenmann 60

145. Heads and Tails: Notes on Sauropod Dinosaurs.

Holland 16

146. Dipterus Remains from Upper Devonian of

Colorado. Eastman .10

147. Notes on Tropical American Tettigonoidea (Lo-

custodea). Bruner 1,66

148. New Species of Tortoise from Jurassic of

Utah. Gilmore 15

149. Fauna of Upper Devonian in Montana.

Haynes .46

150. New Sphagebranchus from Bahamas. Eigen-

mann .07

151. Marine Fishes from Colombia and Ecuador.

Wilson 16

152. Apareiodon, a New Genus of Characid Fishes.

Eigenmann 18

153. New and Rare Fishes from S. American Rivers.

Eigenmann 26

154. Three New Species of Characid Fishes. Eigen-

mann & Henn 12

155. The Species of Salminus. Eigenmann 7

• 156. South American Pceciliid Fishes. Henn 40

157. A New Species of Apatosaurus. Holland .... .7

158. Birds of the Isle of Pines. Todd .70

159. Reptiles and Amphibians of the Isle of Pines.

Barbour 20

160. Land and Fresh Water Shells of the Isle of

Pines. Henderson 20

161. Pelecypoda of the Chazy. Raymond 45

162. S. American Crickets, Gryllotalpoidea, and Ache-

toidea, Bruner 1.30

163. Preliminary Catalog of N. American Sphseri-

idse. Sterki .76

164. Directions for Collecting Sphaeriidae and

Aquatic Gastropods. Sterki .10

165. Lepidoptera of the Isle cf Pines. Holland 60

166. Odonata of the Isle of Pines. Kahl .16

167. A Trip to Islands in Lake Brie. Goodrich .... .16

168. Land-Shells of Islands at Western End of Lake

Erie, etc. Clapp 20

169. Orthoptera of the Isle of Pines. Holland &

Kahl 15

170. Obituary Notes. G. A. Link, T, A. Mills, Boyd

Crumrine, E. M. Bigelow, Holland ...... .10

171. Two New W. African Rhopalocera. Holland. $0.16

172. Botany of the Isle of Pines. Jennings...... 2.00

173. List of Hymenoptera of Isle of Pines. Holland .10

174. Some Species of Farlowella, Eigenmann 80

175. List of S. American Lizards, Griffin ,35

176. Leptodeira albofusca (Lac6pede). Griffin.... .10

177. List of Coleoptera collected on Isle of Pines.

Holland & Schwarz 15

178. Rhynchota of the Isle of Pines. Heidemann

& Osborn lo

179. Mammals of the Isle of Pines. Holland 05

180. Report upon Fossil Material Collected in Cave

in the Isle of Pines. Peterson 10

181. New Species of Fern (Polystichum jenningsi).

Hopkins .10

182. Notes upon the Genus Leucophenga Mik, with

Descriptions of New Species. Kahl ...... .30

183. Some Species of Rhamdia, a Genus of S.

American Siluridse, Eigenmann & Eisher. .16

184. New and Rare Species of South American

Siluridse. Eigenmann 30

185. List of the Hypopthalmidae, Diplomystidae, and

some Unrecorded Species of Siluridse. Eisher. .30

186. A Ssmopsis of the Saurian Genus Prionodac-

tylus. Griffin .05

187. Notes on a Collection of Fishes from Ceylon

with Descriptions of New Species. Jordan
& Starks 35

188. Collection of Fishes from Port Said, Egypt.

Jordan & Hubbs ....'. 20

189. A Fossil-bearing Alluvial Deposit in Saltville

Valley, Virginia. Peterson .10

190. Catalog of Collection of Watches belonging

to H. J. Heinz, Stewart, Holland, Cogges-
HALL Payer, 30 cts.; hoards .50

191. Contributions to the Natural History of the

Isle of Pines: Reprinted from Annals, Vols,
VIII-XI, 560 pp., 34 plates, index. Bound
in cloth 5.00

192. Material Discovered in the Upper Eocene of

the Uinta Basin by Earl Douglass and O. A.
Peterson., Peterson 2.25

193. The Fresh-Water Fishes of the Island of For-

mosa. Masamitsu Oshima 2.75

194. On Elephenor, a New Genus of Fishes from

Japan. David Starr Jordan .50

195. A Description of Cypripedium passerinum.

Jennings 10

196. Obituary of Charles Rochester Eastman 10

197. Obituary of Andrew Arnold Lambing 10

198. Obituary of Herbert Huntington Smith 10

199. Obituary of Henry John Heinz 10

(