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HARVARD UNIVERSITY

eral

LIBRARY

OF THE

Museum of Comparative Zoology

JOHNSONIA

Published by
THe DeparTMENT oF MOLLUSKS
Museum of Comparative Zoédlogy
Harvard University

MONOGRAPHS OF THE MARINE MOLLUSKS
OF THE WESTERN ATLANTIC

VOLUME III
Numbers 33-39

Edited by
WILLIAM J. CLENCH
Curator of Mollusks

Museum of Comparative Zoélogy

PRINTED AT
BOTANICAL MUSEUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS
1954-1959

Henry A. PIcssry,

MUS. COMP. Z08L
LIBRARY

JUN 261959

HARVARD
UNIVERSITY

[ii]

1889

Henry A. Piuspry, apout 1900

HENRY A. PILSBRY
December 7, 1862—October 26, 1957

T is our pleasure to dedicate this third volume of Johnsonia to Dr. Pilsbry. Though
he is perhaps most famous for his work on land mollusks, particularly those of North
America, and his revision of the helices, he published many papers on marine mollusks.
In fact, his publications in the latter field, though a small part of his total studies,
probably exceed those which many workers produce in the course of a lifetime. His
main studies on marine mollusks were those done in the first series of the Manual of

[ iii

Conchology which he took over completely beginning with volume 10. The entire series
consisted of 17 volumes and covered the cephalopods, chitons, scaphopods and marine
gastropods of the world. The volumes on chitons and scaphopods are particularly fine
and are classics in the field of malacology. In addition to the Manual, Pilsbry published
many separate papers on marine shells, outstanding among these being his Catalogue of
the Marine Mollusks of Japan.

Henry A. Pilsbry was born on a farm near Iowa City, Iowa on December 7, 1862.
He was educated in the local public schools and the University of Iowa from which he
received the degree of Bachelor of Science. In 1900 he received an honorary ScD from
the University of Iowa and in 1940 a second honorary degree was conferred upon him
by the University of Pennsylvania. Though he worked for a short time for newspapers,
publishing houses and as librarian at the Davenport Academy of Sciences, his spare time
was always spent in collecting and studying mollusks. His first paper, published in 1882,
was on the land and freshwater shells of Iowa City and until he went to the Academy
of Natural Sciences, Philadelphia where he inherited the work on the marine series of
the Manual of Conchology his interest was entirely in land and freshwater shells.

Dr. Pilsbry joined the staff of the Academy in December 1887 as assistant to George
W. Tryon, Jr. With the death of Mr. Tryon the following year, Henry Pilsbry, as a
young man of 26, became editor of the two series of the Manual of Conchology and Con-
servator of the Conchological Section of the Academy. Later, he was given the title of
Curator of the Department of Mollusks and Marine Invertebrates. In 1889, he became
editor of the Nauti/us and there are numerous papers on marine mollusks by him in this
journal as well as in the publications of the Academy.

Pilsbry’s interest in marine animals was not confined to mollusks for he published on
brachiopods and barnacles and became the world authority on the latter. He published
many papers on this group, the first appearing in 1896; his monograph on the sessile
barnacles which appeared in 1916 is still a basic work in this field. Interest in the cirri-
peds continued throughout his life and at the time of his death he was working on a
paper on barnacles.

The continuation of his early interest in paleontology is evidenced by the many studies
he made of fossil faunas, many of which were based on marine horizons. He was author of
the sections on scaphopods, chitons and gastropods for Zittel’s Textbook of Paleontology.

A brief account of Pilsbry’s life and a complete list of his papers up through Decem-
ber 1939 was compiled by Dr. H. B. Baker and published by the American Malacologi-
cal Union in 1940. Following Pilsbry’s death, Baker, who had been his associate and
close personal friend for many years, wrote a fine, very human and understanding account
of the life of this remarkable man (Nautilus 71, no. 3, pp. 73-83, January 1958). In
this same volume of the Nautilus, which was devoted entirely to Pilsbry, there are ac-
counts of his work on Marine Malacology by Jeanne S. Schwengel (pp. 87-89), on Fossil
and Marine Mollusca by Axel A. Olsson (pp. 89-94), and his scientific contributions
made from 1940 to 1957 were given on pages 104-112.

All branches of Malacology owe a great debt to this man whose keen understanding
of species and their relationships has brought order out of chaos in many groups.

—R. ID. TurNER

[ iv ]

CONTRIBUTORS

Merrill E. Champion ; . #989
William J. Clench : ; , . 85% 365-39
Robert Robertson __. ' : , : : . ST ae
Joseph Rosewater. ; ; : : . 88
Ruth D. Turner : ; ; ; , 33; 384; 35; 86; 38; 389

ACKNOWLEDGEMENTS

We are grateful to a large number of persons who have donated or loaned much of the
material upon which our studies were made. All photographs, unless otherwise noted,
were made by Frank White, staff photographer for the Biological Laboratories, Harvard
University.

STAFF

Editor Assistant Editor
WILLIAM J. CLENCH MerrRILL E. CHAMPION

Associate Editors
R. Tucker Aspott JosEPH C. BEQUAERT JULIA V. CLENCH Ruta D. TuRNER

Business Manager
Ruru D. Turner

bed

CONTENTS
Figures refer to pages

GENERA AND SUBGENERA

Amaea, 344
Anchomasa, 22
Atrina, 310

Bailya, 243

Barnea, 19, 68, 159
Cabestana, 200
Chaceia, 66
Charonia, 198
Columbarium, 330

Conus, 829

Cymatium, 197, 231, 242

Cymatriton, 210
Cyrtopleura, 34, 41
Diodora, 842
Diplothyra, 118
Distorsio, 235

Kpitonium, 844

Gabrielona, 257
Gutturnium, 224
Hatasia, 92
Hemitfusus, 187
Hemitoma, 3385
Jouannetia, 133, 188
Lignopholas, 98, 188
Linatella, 198
Martesia, 101
Melongena, 165
Monoplex, 227
Murex, 33
Nettastomella, 141
Parapholas, 128
Particoma, 114
Penitella, 70, 158

Phasianella, 255, 280

Pholadidea, 89, 159
Pholadopsis, 136
Pholas, 44

Pinna, 301
Pugilina, 184
Ranularia, 204
Rexmela, 170

Rhysema, 236

~ Scobinopholas, 35

Sconsia, 829
Scyphomya, 159
Septa, 214
Servatrina, 314
Thovana, 48
Tricolia, 260
Xylophaga, 145

Zirfaea, 54

FAMILIES COMPLETED FOR THE WESTERN ATLANTIC

Phasianellidae

Cymatiidae Melongenidae

Pholadidae Pinnidae

prvi

HISTORICAL REVIEWS

The Works of G. E. Rumphius, p. 826 The Bolten Catalogue, 1798, p. 283

BOOK REVIEWS
Olsson, A. A., A. Harbison, W. G. Fargo and H. A. Pilsbry, 1953, p. 328
Carcelles, A. R., 1953, p. 64 La Rocque, A., 1953, p. 64

Abbott, R. T., 1954, p. 63

NEW SUBFAMILIES, GENERA, SUBGENERA, SPECIES,
SUBSPECIES AND NAMES

Tricolia a. beaui, 265 Penitella fitchi, 71
Columbarium brayi, 330 Melongena c. johnstonei, 178
Cymatium caribbaeum, 204 Lignopholas, 98
Chaceia, 66 Cymatium r. occidentale, 214
Lignopholas clappi, 99 Tricolia a. pterocladica, 264
Tricolia a. cruenta, 267 Rhysema, 236
Cymatriton, 210 Tricolia thalassicola, 271

== oe + k

ERRATA

No. 34, p. 102, last name in key should read Particoma

No. 34, p. 145, Xyloghagus Meuschen should read Yylophagus

[ vii |

A Partial Analysis of the Molluscan Fauna
of the Western Atlantic

In the table below, we list only the number of species and their synonyms which have
been considered in Johnsonia for the Western Atlantic. Species included in Johnsonia,
but not known to occur in the Western Atlantic, have been excluded from this tabulation.

No single check list has ever been attempted to cover the entire Western Atlantic.
Regional check lists have appeared within the past several years, such as Johnson’s Ma-
rine Mollusca of the Atlantic Coast from Labrador to Texas, several papers by Carcelles
covering Argentina, and a check list by Lange de Morretes for Brasil. To collate these
reports and many others which have been published covering much smaller geographic
areas would still leave rather large gaps in many coastal regions between Greenland and
southern Argentina.

A question frequently asked is ‘‘How many species of mollusks are to be found in the
Western Atlantic?’’ We don’t know, of course, but we can make an estimate that the
molluscan fauna of this area would exceed 6,000 species and subspecies, but probably
would not reach 8,000 species and subspecies.

To date, in Johnsonia, 488 species and subspecies from this region have been mono-
graphed. Though only a fragment of the total number it is sufficiently large to justify a
report upon this number of recognized species and their synonyms. The ratio between
valid species and the synonyms involved will change with each volume completed. The
over-all ratio, however, should remain within fairly narrow limits because the groups
covered have included rare as well as common species from a wide variety of habitats
belonging to both the bivalves and the gastropods.

The synonyms listed below are those which have been described as ‘‘species.’’ Mis-
spellings of names or misidentifications have not been considered as synonyms in this
tabulation.

Johnsonia No. of species No. of synonyms ratio
Vol. 1 210 338 1.61
Vol. 2 169 247 1.38
Vol. 3 59 141 2.64

Total 438 species 726 synonyms _ _1.64

Many regional distribution patterns are beginning to show in our locality records. We
have not, as yet, covered enough of the commoner species to be at all certain about the
limits of many smaller geographic areas. There are still long stretches of coast line,
particularly in Central and South America, from which there is little or no material
available in our various museums.

Pages I-VIII published June 26, 1959

[ viii ]

Mls. COMP. ZOOL. |

LIBRARY
JOHNSONIA MAY 18 1954
7 Published by HARVARD
THE DEPARTMENT OF MOLLUSKS UNCTERSITY

Museum of Comparative Zoélogy, Harvard University

Cambridge, Massachusetts

MAY 17, 1954 PHOLADIDAE VOL. 3, NO. 33*

RHE FAMILY PHOLADIDAE IN THE
WESTERN ATLANTIC AND THE EASTERN PACIFIC
PART I— PHOLADINAE
BY
Ruta. D. Turner

The Pholadidae are a family of highly specialized bivalve mollusks adapted for boring
into wood, soft rock, shells, peat, hard clay and mud. They are characterized by having
accessory plates in addition to the normal bivalve shell, and for this reason were early
classified with the barnacles and the chitons in the Multivalva.

Economic IMPORTANCE

As has been pointed out previously (Clench and Turner, 1946), the financial loss due
to damage caused by wood-boring mollusks probably exceeds by far the amount realized
from the use of mollusks for food and all other purposes. Though the entire family ‘Tere-
dinidae is exceedingly destructive, only a few genera of the Pholadidae are of importance
in this respect.

Martesia is the most destructive genus in the family as all species in the genus are wood-
borers and all may do considerable damage to water front structures. Martesia striata
Linné is the best known and most destructive species and in some localities such as Cavite,
Luzon, Philippine Islands, it may be even more destructive than the Teredinidae. This

Plate 1. Zirfaea crispata Linné, Rye Harbor, New Hampshire (about 2x). Photographed in a test tube un-
der water to show the extended siphons; the external anterior adductor muscle, which is covered only by
a thin layer of periostracum; the small mesoplax and the expanded foot.

* Johnsonia, Volume 38, starts with Number 33.

2 JOHNSONIA, No. 33 Pholadidae

species has been carried nearly all over the world by man and _ has recently been intro-
duced into Sydney Harbor, Australia where it is doing considerable damage. It is also a
problem in San Juan, Puerto Rico and Pearl Harbor, Honolulu.

Species in the genera Xylophaga and Martesia, though generally wood-borers, have
been known to attack underwater cables causing short circuits (Bartsch and Rehder, 1945
and Purchon, 1941). Species in the genus Xylophaga are usually found in floating or
waterlogged wood. They seldom occur in fixed structures, but an occasional attack may
‘ause considerable damage. They may also be found in considerable numbers in buoy
markers and lobster pot floats.

Of the rock-borers, only Penitella penita Conrad has been recorded as definitely de-
structive. At the time of the San Francisco Bay Marine Piling Investigation, it was
found that this species had penetrated the concrete jackets of the piling in several local-
ities about Los Angeles, California. They were boring from a height of two feet above
mean low water down to the mudline. It was found, however, that they occurred in
numbers only in inferior grades of cement.

Diplothyra smithi Tryon and Penitella conradi Valenciennes are both shell-borers, the
former going chiefly into Ostrea and the latter into Ha/liotis. Neither appears to do any
real harm to its host, though heavily infested shells of Hadiotis are, of course, useless in
the jewelry trade.

Most species of the Pholadidae are not of any real economic importance, except per-
haps as their borings may aid in minor changes of the coast line and in the reduction of
wood in the sea. A few species are used as food occasionally though none is of commer-
cial value. Most species are too difficult to remove from their burrows and too full of
sand to make them worth the effort. Pholas dactylus Linné was used for food by the
early Romans and according to Jeffreys (1865) it was used both for food and fish bait in
Normandy and Dieppe. Cyrtopleura costata Linné was at one time sold in the markets in
Habana, Cuba but it is too rare to be of commercial value today. Mr. E. P. Chace of Cali-
fornia writes that the ‘“‘wart-necked piddock’’ makes an excellent chowder—one specimen
being sufficient for two people—but the work of digging out the specimen is considerable.

Novres oN EcotoGy anb DIstTrRiIBuTION

The family Pholadidae is world wide in distribution with most species occurring from
the intertidal zone to a depth of perhaps 250 fathoms. Much greater depths have been
recorded for some species but it is uncertain whether these specimens were taken dead
or alive. A few species appear to be restricted mainly to floating wood and may be con-
sidered pelagic. ‘The family as a whole is restricted locally by the specialized habitat re-
quired foreach group. Species in the genus Zfaea prefer salt marsh peat and stiff mud,
while Penitella and Parapholas bore into much harder shales and sandstones and Pholas
-dactylus Linné may attack gneiss. Martesia and Xylophaga, as borers, are restricted
mainly to wood.

The greatest development of the family appears to be inthe Eastern Pacific which has
13 genera including 23 species, while in the Western Atlantic there are only 8 genera
with 18 species. There are seldom more than two or three species per genus in a given
region and often there is only one. For the genera, Zirfaea, Barnea and Pholas, there
are Companion species in the Western Atlantic and the Eastern Pacific and it would ap-
pear that in the geologic past they were probably one species. The genera Penitella,

Western Atlantic JOHNSONIA, No. 33 3

Parapholas, Pholadidea and Chaceia’ are not found living in the Western Atlantic so far
as is known, but an internal cast of an Eocene (/) fossil taken at Niquero, Oriente, Cuba,
would indicate that a representative, probably of the genus Penitella, was living in the
West Indies at that time. Whitefield (1885) reports a Parapholas from the Eocene marls
of New Jersey and von Ihering (1907, p. 830) reports a species from the Eocene of Pata-
gonia which he refers to Martesia but which on the basis of his figures probably belongs
in the genus Pholadidea. ‘These genera are now restricted to the Pacific with the excep-
tion of Pholadidea, one species of which is found in Europe. Only Zirfaea crispata Linné
and Barnea truncata Say are shared by the Western Atlantic and the Eastern Atlantic
and it would appear that B. truncata Say is probably a fairly recent introduction into
West Africa. Martesia striata Linné, like many species of Teredo and Bankia, is often
found in European waters, having been carried there in flotsam by the Gulf Stream, but
it does not appear to breed in these waters.

The genus Zirfaea is restricted to the colder portions of the temperate and the boreal
regions of the northern hemisphere and the genus VJ'a/ona is restricted to Africa. All
other genera in the family are found both north and south of the equator with the great-
est abundance in the subtropical and warm temperate regions. The occurrence of several
northern species in Lower California may be explained by the upwelling of cold water
at several points along the coast, according to a letter received from J. E. Fitch of the
California Fisheries Laboratory, who has also kindly sent us material from that region.

A detailed account of the ecology of Zirfaea gabbi Tryon (= Zirfaea pilsbryi Lowe) is
given by MacGinitie (1935), and J. E. Fitch (1953) gives a brief account of the habitat

Plate 2. Diplothyra smithii Tryon. A specimen of Crassostrea virginica Linné split lengthwise showing a heavy

infestation of D. smithii Tryon. Chesapeake Bay, Maryland (about 14x).
MUG. CGMP. Z00L.

‘This new genus will be described in the second part of this study.

4 JOHNSONIA, No. 33 Pholadidae

and distribution of five species of California pholads. Scattered observations by numerous
authors such as Dall, Verrill, Morse and others are noted under the species concerned.

Nores ON THE Lire Hisrory oF THE PHOLADIDAE

Very little is known concerning the life history of the various species of this family.
No one species has been completely studied, but varying phases of several species have
been worked out and reported upon. Sigertoos (1894) worked out the early embryology

Plate 3. Zirfaea crispala Linné (young spat).' Fig. 1. At the time of settlement. Fig. 2. With first row of
imbrications which are produced immediately after settlement. Fig. 3. With first row of imbrications well
developed and showing the beginning of the elongation of the posterior slope and the production of the
apophyses (all about 100x).

' All figures from C. M. Sullivan 1948, Bivalve Larvae of Malpeque Bay, P.E.I., Fisheries Research Board
of Canada, Bulletin 77. Weare grateful to Dr. J. C. Medcof and Dr. C. M. Sullivan for the loan of the nega-
tives for these prints.

Western Atlantic JOHNSONIA, No. 33 5

of Barnea truncata Say which he studied at Beaufort, North Carolina. In this species
the sexes are separate, fertilization is external and the eggs and sperm are extruded into
the water, usually during a few weeks in late April and early May. The eggs are small
and rather transparent. Development is so rapid that on warm days the larvae are free-
swimming within three hours after fertilization.

According to Bouchard-Chantereaux (1879) fertilization in Zirfaea is internal and the
young are ejected from the mantle cavity of the adult in the early veliger stage. At
Malpeque Bay, Prince Edward Island, Sullivan (1948) found the mature veliger larvae
of Zirfaea crispata Linné in plankton hauls, from the middle of June to mid-July, when
the temperature of the water ranged from 15° to 22° C. She described and figured the
larvae and the young spat. Her figures, which we reproduce, show that the apophyses do
not develop until the time of settlement. The embryonic shell is about 0.14 mm. in
length and nearly circular in outline, with a straight hinge line. Thorson (1946) also il-
lustrates the veliger larvae of this species and he states that in the Sound off Helsingor,
Denmark, the larvae are found in the plankton from June through February with the
greatest abundance in September.

It has been shown experimentally by Thorson (1946) that the larvae of Zirfaea cris-
pata Linné, like those of many marine bottom invertebrates, are able to select actively
the substratum on which they settle. He states that in experimental bottles ‘‘metamor-
phosing stages of Zirfaea crispata Linné were found in numbers boring into cork floats
and only there.’’ He believes that probably most pelagic larvae can delay metamorpho-
sis for a short period of time if they are not on a suitable substratum and he further con-
cludes that even if the current on the bottom is slow this delay would allow for increased
survival and dispersal. Though neither Dr. Sullivan nor Dr. Thorson mentioned the at-
tachment of the young spat by a single byssus thread at the time of settlement, such is
probably the case. The production of a single byssus in the Teredinidae is well known
and our observations would seem to bear this out for Zirfaea. Newly settled spat in a
test-board submerged at Beverly, Massachusetts were observed to have constructed a
small conical covering of cemented wood scrapings for protection during the last stages
of metamorphosis. This has also been observed in the Teredinidae, and at this time the
young shipworms attach themselves by a single byssus thread, probably to prevent being
moved about by the currents. It was also observed on this test-board that the young Z.

1/2 mm.

Plate 4. Prodissoconch of Xylophaga atlantica Richards. Fig. 1. External view of valve showing marked con-
centric growth lines. Fig. 2. Internal view of valve showing the two pronounced hinge teeth. Fig. 3.
Specimen in the late veliger stage showing the beginning of the elongation of the shell. All specimens
taken from the mantle cavity of a preserved specimen which had been taken by the Albatross at station
2550, about 160 miles east of Barnegat Bay, New Jersey.

6 JOHNSONIA, No. 33 Pholadidae

crispata settled only on the horizontal upper surface of the board. Though Zi-faea usu-
ally are not wood-borers, they are occasionally found in waterlogged wood but their
presence in a test-board is most unusual.

On the basis of material which has been available for study it appears that fertilization
in Xylophaga dorsalis Richards is internal and the young are retained within the parent
until the late veliger stage and time for settlement. The accompanying plate shows the
shell of the young taken from a preserved adult specimen. Thus it would appear that in
the Pholadidae as in the Teredinidae ovoviviparity is at best of no more than generic
value taxonomically.

Moore (1947) states that the adults of Martesia striata Linné are alternating herma-
phrodites and Purchon (1941) records the same for Xylophaga dorsalis Turton. In addi-
tion Purchon discusses the presence of an accessory genital organ in Xylophaga and the
storage of spermatozoa in an organ which he terms a ‘‘vesicula seminalis. ** This he believes
would allow for self-fertilization, a condition which would be of distinct advantage for a
species living in isolated colonies, usually in floating or waterlogged wood as is the case
with Xylophaga.

The rate of growth undoubtedly varies greatly in all members of the Pholadidae, though
we have specific data only on such economic species as Martesia striata Linné and Mar-
tesia cuneiformis Say. Once established in their burrows, the rate of growth depends
largely upon the hardness of the substratum and the amount of crowding. 'Test-board rec-
ords show that specimens may produce a callum in less than a month and when the shell
is only 4 mm. in length. Such specimens, generally termed stenomorphs, may continue
to live for some time and are capable of producing large numbers of eggs. Under most
favorable conditions a specimen may reach 45 mm. in length before producing a callum
and may remain in an active state for several months. Specimens from test-boards at San
Juan, Puerto Rico had reached a length of 40 mm. in six months and at Bahia, Brasil,
specimens of Martesia cuneiformis Say taken from a board submerged on May 8, 1945
and removed on June 2, 1945 had reached a length of 11.5 mm.

The average length of life for any one species of pholad, the breeding age or the num-
ber of eggs produced, and the duration of the free-swimming larval life are questions
still unanswered.

Additional notes on life history and habits, if known, are included in ‘‘Remarks’’ under
the individual species.

Meruops or Borinc

Since early times scientists have argued over the method by which the various mem-
bers of the Pholadidae bore. Species of the genera Pholas, Parapholas, Penitella, Di-
plothyra and Zirfaea bore into substrata varying in hardness from that of stiff clay to
limestone, shale, gneiss and shells such as Halotis and Ostrea. Interesting and rather
amusing accounts of the various theories of the early 1800's are given by G. Johnston
(1850), Forbes and Hanley (1853) and J. G. Jeffreys (1865). Included in their accounts
are the arguments supporting theories that boring was accomplished by means of an acid,
that the foot did the boring using imbedded siliceous spicules and that the shell was the
tool. Buonanni (1684) was the first to believe that the shell was the instrument of boring
and Osler (1864), hoping to settle the argument between those who thought boring was
accomplished by acid and those who believed in the mechanical theory, spent considera-

Western Atlantic JOHNSONIA, No. 33 +i

ble time observing living animals. He presented a rather detailed paper on his views that
boring was mechanical, the shell being used as a tool. The arguments continued, how-
ever, and Cailliaud (1855-1857), one of the most ardent of the researchers in this field,
wrote numerous papers on the methods of perforation of rocks by Pholas. Dubois (1892,
p. 84) believed that the siphons alone were responsible for the boring, stating that the
musculature of the siphons was sufficient to turn the animal completely around in its
burrow. This would mean that the animal was boring like an auger. The action of the
foot with the use of sand grains is the theory put forth by Smith (1894) but this could
not explain the wood-borers. A rather comprehensive account of the boring mechanisms
of several forms that occur in Puget Sound is given by F. E. Lloyd (1887). He goes into
particular detail on the musculature of P. penita Conrad, showing that in forms which
have a ventral adductor muscle, the anterior adductor muscle is divided into two parts.
The anterior portion, which he terms the accessory anterior adductor, opposes the pos-
terior adductor muscle. The contraction of the accessory anterior adductor muscle closes
the valves anteriorly and opens them posteriorly. ‘The posterior portion, which he terms
the anterior adductor muscle, opposes the ventral adductor muscle. The contraction of
this portion of the muscle spreads the valves apart ventrally and brings them in contact
with the sides of the burrow.

The belief that an acid isemployed in boring is no longer held today, but many of the
other ideas set forth by early workers fit into the complex picture. The summation of
all of these theories along with observations made in the laboratory would indicate that
the task of boring is not accomplished by any one organ alone. ‘The shell may be con-
sidered the tool which is manipulated by the adductor muscles and is aided by the foot,
siphons, mantle, water, and accessory sand grains. An exact procedure does not hold
true for all members of the group and even appears to vary somewhat within the species
depending upon the hardness and compactness of the substratum into which the speci-
men is boring. MacGinitie (1985) has given an excellent account of the boring of

Plate 5. Penitella penita Conrad. A typical snug burrow of a Penite//a boring in soft shale showing the depth
to which this group usually bores. White’s Point, San Pedro, California (about 13x).

8 JOHNSONIA, No. 33 Pholadidae

Zirfaea gabbi Tryon (= Zirfaea pilsbryi Lowe) and our observations on Zifaea crispata
Linné agree with his. Active specimens of this species, of Barnea truncata Say, and of
Martesia striata Linné, which were observed in the laboratory, all followed the same
general pattern. The foot was first attached to the anterior end of the burrow, the con-
traction of the anterior portion of the anterior adductor muscle and the ventral adductor
muscle closed the valves anteriorly as much as possible, and the shell was brought as far
forward in the burrow as possible. Then the forcible contraction of the posterior adductor
muscle and the posterior portion of the anterior adductor muscle, by opening the valves
ventrally and closing them posteriorly, brought the recurved imbrications of the beaks
forcibly against the sides of the burrow. The insertion of the anterior adductor muscle
on the umbonal reflection in an external position, allows the two sets of muscles to act in
opposition to each other and produce a rocking and heaving motion of the valves with the
umbos acting as a center.

Mechanically the shell is only capable of boring into a substratum that is as soft as the
shell itself or softer. As the shells are composed of aragonite it is possible for them to
bore into soft limestone without difficulty. When they are boring into a harder rock such
as gneiss, there must be some elements in the rock which are softer than the shell itself
or else the rock must be of a friable nature. In such cases the imbrications of the shell
‘an work out the softer portions of the rock, such as the mica, and the harder particles
of quartz, feldspar and garnets then fall out. Many of these harder particles apparently
are caught in the mucus around the foot, on the thickened frontal margin of the mantle,
and many have been observed impressed in the cephalic hood of Zfaea. Particles are also
‘aught between the rows of imbrications on the anterior portion of the valves and these
may be in sufficient quantity to cover the worn imbrications. ‘The result is that in species
such as Pholas dactylus Linné the stone in many cases is worn away with particles of the
same hardness.

While watching Barnea truncata Say and Zirfaea crispata Linné in an aquarium, it
was noticed that they are capable of producing a strong and sizeable stream of water just
ventral to the foot. This jet is forced out of the mantle cavity, the mantle edge being
held close to the foot except for a small area just ventral to the foot. The stream is di-
rected forward and probably is used to flush out the anterior end of the burrow.

Some species, such as Zirfaea pilsbryi Lowe (MacGinitie 1935) and Cyrtopleura costata
Linné, excavate deep, rather roomy burrows and are capable of moving up and down in
them at will. During high tide when they are feeding, the siphons may be seen extending
a considerable distance out of the opening. Other species such as Penitella penita Conrad,
as shown in the accompanying plate, excavate a very snug burrow in which movement
is limited. This seems to be true of most boring into a hard substratum. The length of
the burrow varies greatly with the different species. Some make burrows which are just
about equal to the length of the shell, as is the case with Martesia striata Linné, while
others may be several times the length of the shell. The type of burrow, however, is
quite consistent for each species.

Specimens of Barnea truncata Say, Zirfaea crispata Linné and Martesia striata Linné
when removed from their burrows are completely incapable of re-burying themselves. If,
however, a shallow depression is made in the substratum just sufficient for the animal to
attach its foot and bring the anterior edges of the shell into play, it will readily excavate
a new burrow.

Western Atlantic JOHNSONIA, No. 33 9

Little is known as to just how the young, particularly of the rock-boring forms, get
started. One of the quaint early theories was that the parent could produce an acid in the
tips of the siphons and with this make holes in the rock for the reception of the young.
From observations made on Zirfaea crispata Linné it would appear that the young prob-
ably attach themselves by a single byssus thread in the protection of a minute crevice
while metamorphosis takes place and the shell changes into a boring tool.

Nores ON THE ANATOMY AND PHYSIOLOGY

The shell is the most unique anatomical feature of the Pholadidae, as no other family
among the Pelecypoda has developed accessory plates. In addition, only the Teredinidae
share with the Pholadidae the apophyses and the externally placed anterior adductor
muscles. While no over-all study of the anatomy of the Pholadidae has been made, the
gross morphology of several species has been reported upon and the detailed anatomy of
a few is known. The first work was that of Poli (1791) in which he illustrated the anat-
omy of Pholas dactylus Linné, including detailed drawings of the gills, the circulatory
and digestive systems. Fischer (1858-1860) described the general anatomy of several
widely separated species including Pholadidea melanura Sowerby, Parapholas acuminata
Sowerby and Jouannetia globosa Sowerby. Egger (1887) described in great detail the
anatomy and histology of Jowannetia cumingu Sowerby. In all cases there are no real
basic differences in the soft anatomy between these forms and other bivalves. Striking,
though not basic differences, include the elongate gills of the pholads which extend into
the incurrent siphon and the large triangular to strap-shaped labial palps. Observations
on Nettastomella rostrata Valenciennes show that in this species the outer demibranchs
of the gills are somewhat reduced, and that the labial palps are very small compared with
those found in Zfaea and in other genera. In _Xylophaga only one demibranch remains
and this is considered by Ridewood (1908) to be the inner one while Purchon (1941) be-
lieves that it is the outer one. Purchon also points out that the labial palps of Xylophaga
are small and spindle-shaped, with a greatly reduced sorting mechanism. This is a condi-
tion that would be expected in wood-boring forms usually found living in floating wood,
where most of the material brought in by the siphons is usually more or less uniform in
size. In the mud and rock-boring forms, however, the ciliary sorting mechanism is
highly developed.

The structure and function of the gills, ciliary mechanism, food tracts and method
of feeding of Zirfaea crispata Linné were first described by Alder and Hancock (1851).
Kellogg (1915) has given an excellent account of the ciliary mechanism in several groups
including Barnea, Zirfaea and Penitella. Basically the ciliary mechanism in the species
he examined was similar to that in other bivalves; however, several interesting differences
were noted. Among these was the development of a covered food groove in Zirfaea to
insure the delivery to the palps of the food collected on the gills. Of particular interest
is his description of the ‘‘collecting membrane’’ found in Barnea { = Cyrtopleura] costata
Linné. This membrane, extending from the posterior portion of the visceral mass, col-
lects the excess silt and deposits it in a ball well out in the incurrent siphon whence it is
expelled by a sudden contraction of the posterior adductor muscle. This apparently al-
lows feeding at all times even in the very muddy water where this species often lives.

The crystalline style in all species examined was unusually large and, in the callum-

10 JOHNSONIA, No. 33 Pholadidae

building forms, after the foot has atrophied, the style can be seen in the style sac pro-
truding slightly from the anterior end of the visceral mass. Graham (1949) made a com-
parative study of numerous molluscan stomachs, including several species of pholads and
showed their resemblance to the general type.

DuBois (1892) worked on the anatomy and physiology of Pholas dactylus Linné, but
restricted his studies almost entirely to the siphons, and Purchon (1941) has dealt in de-
tail with the biology and relationships of Yylophaga.

The callum-building pholads present two unique and rather interesting phenomena.
In most species the animal appears to be able to store a considerable amount of calcium
in its system, so that when the time comes the callum is laid down very rapidly. Among
the thousands of specimens examined very few were found with the callum partially
built. This undoubtedly is one of the reasons why the young stages of callum-building
forms have so often been described in separate genera. Early workers, finding adult speci-
mens (ones with a completed callum) which varied greatly in size, believed that the cal-
lum could be reabsorbed, that the animal could continue growth and then produce a
second or even a third callum ata later date. This, however, is no longer considered the
‘ase, as in all of the callum-building forms the foot is absorbed once the callum is com-
pleted and boring has ceased. The muscles which work the valves during boring opera-
tions do not atrophy as might be expected, because a slight movement of the valves is
still possible, and this movement is important for the circulation of water within the
burrow.

The luminous properties of Pholas dactylus Linné have intrigued naturalists since early
Greek times. In his writings Pliny speaks of the luminous fluid as being so abundant
that the hands and mouths of those who ate them shone. An excellent summary of the
experimental work on the bioluminescence of this species is given by Harvey (1952, pp.
255-267). ‘Tomlin (1920) reports that specimens of Yylophaga praestans Smith were
luminescent at night, but Purchon did not report this for XY. dorsalis Turton. So far as
is known no other species of Pholadidae is luminescent, with the possible exception of
Barnea candida Linné which, according to Okada (1927), has a weak momentary lumines-
cence when opened. However, the two species most closely related to P. dactylus Linné
have not been investigated in this respect. The Western Atlantic form, P. campechiensis
Gmelin, occurs mainly below low tide line and is almost unknown except as dead beach
specimens. Pholas chiloensis Molina of the Eastern Pacific is more common and should
be investigated for this property. Living specimens of Zirfaea crispata Linné and Barnea
truncata Say which we observed both in the laboratory and in the field showed no lumi-
nous properties. |

All species of the Pholadidae, so far as known, live under strictly marine or only
slightly brackish water conditions, with the possible exception of Martesia rivicola Sow-
erby, an Indo-Pacific species, which is recorded as occurring in floating wood in fresh
water. Some species, however, appear to be able to withstand dilutions of sea water down
to 50% for short periods of time. This would certainly indicate that, though these spe-
cies are strictly marine, they have the ability to adjust themselves to the hazards of their
intertidal habitat. Unlike Venus, Mytilus and others, they are unable to retract com-
pletely within their shells and so shut out the environment when conditions are adverse.
A heavy rainfall at low tide could quickly fill the burrows of Zirfaea and Barnea with

Western Atlantic JOHNSONIA, No. 33 11

nearly fresh water and consequently the ability to adjust to such changes probably has
some survival value.

Zirfaea crispata Linné appears to be able to withstand a considerable range of tem-
perature. Specimens placed in a jar of sea water and exposed to freezing temperatures
became inactive when slush-ice was formed. On thawing, however, the animals again be-
came active. In the summer they can tolerate water temperatures at least as high as 75° F.

SHELL MorrHo.ocy

The pholad shell is complicated and highly specialized. Besides the two valves common
to all pelecypods, some members of the Pholadidae may have as many as four accessory
plates and all members have at least one. In some (the Pholadinae and Xylophaginae)
there is no basic change from the young to the adult shell, while in others (the Martesii-
nae) there are two distinct stages in shell growth. The young shell is beaked and widely
gaping anteriorly, while in the adult the anterior gape is closed by a calcareous deposit,

protoplax

mesoplax

PUY 4011a4Ss0d

anterior margin
of shel

umbonal-Ventral sulcus

chondrophore umbonal reflection anterior adductor muscle

apophysis

posterior adductor muscle
accessory anterior
adductor muscle

siphonoplax

pallial line

posterior end

anterior margin of shell

so

umbonal-ventral ridge

Hial line
hed pallial “sinus

2 ventral adductor muscle

ventral condyle
Plate 6. Diagrammatic, composite drawing of a pholad shell. Fig. 1. External view of the valve to show the
approximate relative positions of the various accessory plates and the parts of the valve proper. Fig. 2.
Internal view of the valve to show the general arrangement of the muscle scars, the placement of the apo-

physis and other parts.

12 JOHNSONIA, No. 33 Pholadidae

the callum. The young stage which has been termed the ‘‘working or Zirfaea stage’” by
Lloyd (1897) continues as long as the animal is actively boring. During this stage the
shells of several of the genera closely resemble those of Zirfaea; they are beaked anteri-
orly, rounded posteriorly and the mesoplax, the only accessory plate present at that
stage, is a simple transverse plate located across the umbos and beneath the posterior
portion of the anterior adductor muscle. The anterior adductor muscle is protected at
this stage by a chitinous covering, sometimes referred to as the cephalic hood, a term
more commonly used in the Teredinidae. When the excavation is completed and the
animal ceases growing, the pedal gape is closed by a callum, the mesoplax is completed
and the metaplax, hypoplax, and siphonoplax, if present, are developed. This stage,
Lloyd (1897) has called the ‘‘resting stage.’” When the callum develops, the foot atro-
phies and the pedal gape of the mantle closes except for a minute anterior pore. Finally
the completed halves of the callum are joined by a chitinous band, with only a minute
anterior poré remaining open.

Most of the Pholadidae have apophyses, large shelly projections which extend from
beneath the umbos and serve as the attachment area for the large foot muscles and as a
support for the visceral mass. In all of the pholads the dorsal margins of valves are re-
flected anteriorly, so that the anterior adductor muscle is in an external position and
works in opposition to the posterior adductor muscle.

Many of the following terms for the various parts of the shell and the accessory plates
were first introduced by Fischer in 1858.

ProropLax. The term protoplax is restricted to the simple, nearly flat dorsal plate
which is anterior to the umbos and which rests on top of, but does not enclose, the an-
terior adductor muscle. It is essentially the same in young and adult shells and its axis
is longitudinal. It may be in one piece or divided longitudinally into two equal parts.
It may be chitinous or calcareous.

Mesop.ax. The mesoplax is a transverse plate, usually wider than long, which strad-
dles the two valves on their dorsal margin. It protects the posterior portion of the an-
terior adductor muscle and always originates ventrally to the muscle. In the Pholadinae,
the mesoplax is simple and essentially the same in the young as in the adult. In the
Martesiinae, however, the mesoplax in the young is a simple, more or less semicircular
plate situated mainly ventral to the posterior portion of the anterior adductor muscle.
In the adult of these forms it grows dorsally and then anteriorly so as to enclose the pos-
terior portion of the anterior adductor muscle. ‘The mesoplax may be in one or two pieces
and is always calcareous.

Merap.ax. The metaplax is a long, narrow plate which covers the gap between the
two valves on the dorsal margin posterior to the umbos. It is usually caleareous and in
most species is attached to the dorsal margins of the valves by a chitinous fold. The
metaplax may be pointed, rounded or forked posteriorly.

Hyropiax. The hypoplax is the long, narrow ventral plate covering the gape between
the two valves on the ventral margin. It extends from the umbonal-ventral suleus pos-
teriorly, where it may be forked, rounded or pointed. It is usually entirely calcareous
and is joined to the ventral margins of the valves by a chitinous fold.

Western Atlantic JOHNSONIA, No. 33 13

SipHONoPLAX. This term is applied to any structure which is secreted by the mantle
on the posterior margin of the valves, probably for the protection of the siphons. The
siphonoplax may be chitinous or calcareous, the two halves diverging or fused to form a
tube. In Jowannetia the siphonoplax is developed on the right valve only.

CautLuM. This term refers to the closure of the pedal gape. It may be partial or com-
plete, sculptured or irregularly marked. Generally the two halves of the callum do not
quite meet and are joined by a chitinous fold, with only a minute anterior pore remaining
open. In Jouannetia the callum of the left valve is greatly developed and overlaps that
of the right valve. In Nettastomella the calcareous portion of the callum consists only of
a narrow band extending along the anterior margin of the shell; the main portion of the
callum is chitinous. In many species of Martesiinae the callum extends dorsally between
the beaks and covers the anterior portion of the anterior adductor muscle.

UmponaL REFLEcTION. This term refers to the reflection of the dorsal margin of the
valves anterior to and usually over the umbos. It may be closely appressed to the shell
surface, raised well above the surface of the shell, with or without a posterior support, or
the space below the reflection may be septate. The anterior adductor muscle scar usually
covers most of the umbonal reflection.

ApvopuyseEs. These are large, styloid projections, one in each valve, extending from
beneath the umbos to which the foot muscles are attached.

CuonpropHore. This is a modification of the hinge area to support the internal liga-
ment. The chondrophore of the right valve is a small swelling with a central depression,
while that of the left valve is a small shelf-like projection. The chondrophore is found in
only certain groups of the pholads.

SipHONAL Tuse. This term is applied to the tube produced in some members of the
genus Pholadidea. It is composed of agglutinized particles produced as a result of bor-
ing activities. It differs from the ‘chimney’ of the Parapholas in that it is fused to the
siphonoplax.

CuiIMNEy. A tube formed of agglutinized particles produced as a result of boring activ-
ities. It fits over the posterior end of the shell and in some species extends anteriorly
nearly to the mesoplax.

Other specialized terms can readily be understood from the accompanying diagram of
a composite pholad shell on which all the various parts have been labeled. Only in the
genus Parapholas are the three main areas of the shell as clearly marked as in the diagram.
No one genus has all of the accessory parts indicated.

GENERAL REMARKS ON THE CLASSIFICATION

The family Pholadidae is a compact and easily recognized group. All of its genera
possess three or more of several characters (such as accessory plates, callum, umbonal
reflection, external position of the anterior adductor muscle, apophyses, ventral condyle,
and chimney or siphonal tube) found in no other family of Pelecypods except the 'Tere-

14 JOHNSONIA, No. 33 Pholadidae

dinidae. No pholad, however, has the elongate body and pallets of the Teredinidae and
no teredo has accessory plates, callum or chimney.

The main difficulty faced by the early workers in the Pholadidae was the lack of ma-
terial in which all the accessory plates were present. The basic shell in many of the spe-
cies is very similar and when the accessory plates are lacking it is easy to misidentify
them. Inadequately understood life histories resulted in the creation of new genera for
the young stages of callum-building forms. In addition, the species are exceedingly vari-
able. Therefore it is little wonder the early workers, having the extreme forms of a spe-
cies with none of the intermediate ones, described them as different species. It is not
unusual in this family to have adult specimens of a given species ranging from 8 mm. to
45 mm. in length, and the ratio of length to height is nearly as variable. This range of
size and shape is a result of the boring habit of this group. The type of substratum in
which the animals bore and the amount of crowding are largely responsible for the vari-
ation. Species which normally bore into wood become misshapen, small and thick-shelled
when they work in harder materials. Overcrowding produces stenomorphs or greatly
stunted forms which assume all of the adult characters though they are perhaps only 1/10
normal size. Conversely, species such as Penitella conradi Valenciennes {= P. parva
Tryon], which normally bore into Hadliotis or other shells, are equally variable. When
they live in softer material, such as heavy clay, the shell is much larger than normal and
is far more perfectly formed. Consequently in this study an attempt has been made to
obtain large series of preserved material from as many localities as possible in order to
have a good picture of the range of size and shape of the species and the various types of
malformations that normally occur.

On the death of the animal the accessory plates may become detached and isolated and
these on occasion have been described, usually as species of simple gastropods of the type
of Scutum, Acmaea or Patella. Mérch (1876) has pointed out that the dorsal plate of
Pholas (Monothyra) orientalis Gmelin was described in 1779 by J. E. J. Walch as Seutum
dacicum and in 1874 G. and H. Nevill again described it as Seutum ? abnormis. We have
not seen the type specimen of Seutwm abnormis but from the illustration there is no doubt
as to its identity with the oriental species. The apophysis of Cyrtopleura costata Linné
was described by Conrad as Capulus shreevei and it is very possible that Patella acinaces
Lea 1846 is the dorsal plate of Barnea truncata Say. Undoubtedly there are other cases
of such misidentifications which will eventually be clarified; some may possibly have been
described as plates of barnacles. Isolated accessory plates occurring in fossil beds could
prove a real problem to paleontologists.

Though the Pholadidae are known from the Pennsylvanian, the fossil record is meager
and seattered and, of course, none of the soft parts have been preserved. On the basis of
the present record it is impossible to say whether the Pholadidae and the Teredinidae
stemmed from a single ancestor or whether the pholads gave rise to the teredos or vice
versa. There is no question that the two families are closely related and that they either
stemmed from a common ancestor or that one gave rise to the other. There are four dis-
tinct subfamilies in the Pholadidae all of which have accessory plates and an externally
placed anterior adductor muscle. Of these four subfamilies, two have apophyses and two
lack them. In each group there are callum and non-callum producing genera as shown
in the following table.

Western Atlantic JOHNSONIA, No. 33 15

Subfamilies Apophyses Callum Accessory Plates
Pholadinae xX xX
Martesiinae xX X X
Xylophaginae ». 4
Jouannetiinae Ss xX

Family PHOLADIDAE

Shells with a narrow, slit-like to nearly circular pedal gape, which may or may not be
closed by a callum in the adult stage. Anterior portion of the valves imbricate or denti-
culate and often ribbed. This portion is often separated from the posterior portion by an
umbonal-ventral sulcus. Hinge teeth usually lacking, a small chondrophore present in
some forms, ligament if present, internal. Anterior dorsal margin of the valves reflected,
forming the attachment area for the externally placed anterior adductor muscle. The
fused mantle edges in many forms are thickened and muscular. The ventral adductor
muscle, present in some forms, is a thickened muscular portion of the ventral margin of
the mantle. Anterior adductor muscle protected in the young stage by a chitinous cover-
ing often referred to as the cephalic hood. In the adult stage of most forms it is covered
by accessory plates or by a dorsal extension of the callum. The total number of acces-
sory plates in any one species may vary from one in Barneaand Zirfaea to four in Mar-
tesia and Parapholas. These last two forms have a mesoplax, metaplax, hypoplax and
callum. A chimney is produced by Parapholas and Xylophaga. Apophyses are present
in the Pholadinae and Martesiinae, but absent in the Jouannetiinae and XNylophaginae.
For the distribution of the accessory plates in the various genera see the following table.
Pallial sinus usually deeply inset. Siphons united, capable of considerable extension,
smooth or papillose and often enclosed in a chitinous sheath. Siphonal aperture usually
surrounded by cirri. Gills with two demibranchs, except in the XNylophaginae where
there is only one, the outer one according to Purchon (1941). In many groups the gills
are extended into the siphons. Foot well developed, usually truncate and adapted for
suction. The foot in all of the Martesiinae and Jouannetiinae atrophy in the adult stage.

The following arrangement of the genera within the family is an attempt to group sim-
ilar forms from those that are the simplest structurally to those that are the most com-
plex. It is not intended to show any actual or supposed evolutionary sequence. The
entire family is rather old geologically as shown on the following table, which gives the
range in time, so far as is known, of the genera considered in this paper. However, it is
often impossible to place fossil pholads in their proper genus owing to the fact that they
may be only internal casts, that they lack the accessory plates, or that they are young
forms. As represented today the family is composed of numerous well marked genera,
each genus containing few species and seldom are species transitional between any two
genera. This condition is paralleled by other groups in the Mollusca, where families,
which are old geologically speaking, have become differentiated generically but now show
little breakdown on the specific and subspecific levels.

Subfamily PHOLADINAE
Shell not closed anteriorly by a callum in the adult stage. Number of accessory plates
variable but hypoplax and siphonoplax always lacking. Apophyses present. Valves not
divided into two regions by an umbonal-ventral sulcus, except in the genus Zi7faea where

16 JOHNSONIA, No. 33 Pholadidae

the sulcus is rather weak and often barely visible in the adult. Foot well developed, not
atrophying in the adult. Animal not capable of complete retraction within the shell.

Subfamily MarreEslINAE

Shells beaked and gaping anteriorly in the young stage but closed by a callum in the
adult. Number of accessory plates variable, but always lacking a protoplax. Apophyses
present. Valves divided into two distinct areas by an umbonal-ventral sulcus and in ad-
dition, in the genus Parapholas, a ridge extending from the umbos to the posterior ventral
margin divides the valves into three areas. Foot well developed in the young stage but
atrophied in the adult. Animal capable of complete retraction within the shell except in
the genus Chaceia.

Subfamily JoUANNETIINAE

Shell beaked, gaping very widely anteriorly in the young stage and partially to com-
pletely closed by a callum in the adult. The callum in the genus Nettastomella exists as
a peripheral band of calcareous material with the large central portion being chitinous. In

CHART SHOWING THE PROBABLE GEOLOGIC RANGE OF THE GENERA

(The solid lines are not necessarily indicative of a continuous fossil record)

n n
= =
o oO
2 Sg
= 2 > o
ants a> 6 ot he S os
= be a 4 S = ) a
i= = ae S 2 re eee v 0 v = S = »
- x =] — a n mo ~ vo r= © oO Oo I
Se hei ee ee. es eS ee ee oe a <8 tn
= = : 2 3 w x > 2 ro) v 5 = 5 z =
& ao) = rs a q ay es = SS i) of ° ° ‘Oo S)
3 Be = 3 o a = Qo. (|e os Sp RS Oe co)
ee in. ee Sy) et Se eB ee Oe hy Se,
Barnea
Cyrtopleura
Pholas
Zirfaea
P= = || (Clncein
Peasoose ee Ol acne
Penitella
| RParapholas
?----— | Diplothyra
oO —- so Marrtesin
Jouannetia
?-----— =| Nettastomella
Xylophaga

Western Atlantic JOHNSONIA, No. 33 1

the genus Jowannetia the callum is entirely calcareous and greatly produced. Apophyses
lacking, the foot muscles being inserted in the normal position. Accessory plates varia-
ble; a rudimentary mesoplax is present in some species. The siphonoplax heavy, calcare-
ous and in Jouwannetia present only on the right valve. Anterior adductor muscle covered
by the dorsal extension of the callum. Shells divided into two distinct areas by the
umbonal-ventral sulcus. Foot well developed in the young stage but atrophied in the
adult. Animal capable of complete retraction within the shell.

Subfamily NyLoPHAGINAE

Shell beaked and gaping anteriorly throughout life. The beaks are truncated at nearly
right angles, giving the shell a teredo-like appearance. Valves rounded and closed pos-
teriorly, the animal capable of complete retraction within the shell. Anterior portion
sculptured with numerous rows of finely denticulated ridges as in the Teredinidae. Pos-
terior portion sculptured only by growth lines, and separated from the anterior portion
by the umbonal-ventral sulcus. Accessory plates consisting only of a small divided meso-
plax. Callum and apophyses absent. Umbonal-ventral ridge pronounced and usually with
a ventral condyle. Umbonal reflection narrow, simple and closely appressed. Posterior
adductor muscle sear large, generally oval in outline and placed high on the posterior
slope. Foot muscle inserted in the normal position. Foot not atrophying in the adult.

Distribution of accessory parts in the various genera of the Pholadidae

Apophysis Protoplax Mesoplax Metaplax Hypoplax Siphonoplax Callum Chimney
PHOLADINAE

Barnea X X

Cyrtopleura xX X xX

Pholas xX X xX x

Talona X x

Zirfaea xX xX
MARTESIINAE

Chaceia xX xX xX

Penitella x x x x

Pholadidea X xX xX xX

Parapholas xX xX xX xX xX ».4

Diplothyra xX X X xX x

Martesia X x X xX xX
JOUANNETIINAE

Jouannetia xs X X

Nettastomella X xX
XYLOPHAGINAE

Xylophaga D.¢ xX

ACKNOWLEDGMENTS

Many institutions and individuals have been exceedingly helpful in providing material
for this study. Before listing these, I wish to express my appreciation to several members
of the Museum staff for their help: to Dr. William J. Clench under whose immediate
direction the work was done and whose continued interest and philosophy of work have
been most important in helping me solve the many problems concerned in this study;

* Present in some but not all species in the genus.

18 JOHNSONIA, No. 33 Pholadidae

to Dr. Joseph C. Bequaert and Dr. Merrill Champion for their counsel and aid in read-
ing the manuscript; to Professor Henry B. Bigelow and Professor Ernst Mayr for their
many helpful suggestions, and to Professor John H. Welsh of the Biological Labora-
tories for aid on physiological problems.

Several institutions have been very generous in the loan of their material, in locating
types, and in photographing type specimens. For their help I wish to thank the fol-
lowing:

C. G. Aguayo, Museo Poey, Habana, Cuba; Alberto R. Carcelles, Museo Argentino
de Ciencias Naturales, Buenos Aires, Argentina and Juan Parodiz, now on the staff of
the Carnegie Museum at Pittsburgh, Pennsylvania; D. J. Cleghorn, Redpath Museum,
McGill University, Montreal: Ttadashige Habe, Zoological Institute and Seto Marine
Biological Laboratory, Kyoto University, Japan: Leo B. Hertlein, California Academy
of Sciences, San Francisco, California; Myra Keen, Stanford University, Stanford Cal-
ifornia; J. Knudsen, Universitetets Zoologiske Museum, K6benhavn, Denmark; J. C.
Medcoft, Atlantic Biological Station, St. Andrews, New Brunswick; Th. Monod, Insti-
tut Frangais d’ Afrique Noire, Dakar, Sénégal; George Moore, University of New Hamp-
shire, Durham, New Hampshire; Hilary Moore, Marine Laboratory, University of
Miami, Florida: Katherine Van Winkle Palmer, Paleontological Research Institute,
Ithaca, New York; Henry A. Pilsbry, Academy of Natural Sciences, Philadelphia;
R. D. Purchon, University of Malaya, Singapore; Harald A. Rehder and R. ‘Tucker
Abbott, United States National Museum, Washington, D.C.; Emma B. Richardson,
Charleston Museum, Charleston, South Carolina; F. S. Russell, Marine Biological Lab-
oratory, Plymouth, England; 'T. Soot-Ryen, ‘Tromsé6 Museum, ‘Troms6, Norway: C.J.
Trist, Queensland Forest Service, Brisbane, Australia; Tera van Benthem Jutting
vander Feen, Zoologisch Museum, Amsterdam, Netherlands: Henry vander Schalie,
Museum of Zoology, University of Michigan, Ann Arbor, Michigan; W. J. Rees and
G. Wilkins, British Museum (Natural History) London, England; C. M. Yonge, Uni-
versity of Glasgow, Scotland.

From the late William F. Clapp and through the kindness of A. P. Richards and
Dorothy Brown of the Clapp Laboratories, Duxbury, Massachusetts, we received for
study the entire collection of Martesia which they had brought together. This is a par-
ticularly important collection, for all specimens were taken from test-boards, thus estab-
lishing the occurrence and breeding of the species in a given locality, as well as adding
important data on growth rates.

Several friends made a special effort to collect material for which I wish to express my
gratitude. Emery P. Chace of Lomita, California, sent not only his collection but also
preserved specimens he had collected in and around San Pedro, California. Ruth Coats
sent a large collection of preserved material from Tillamook, Oregon. Special thanks are
due John Fitch of the Terminal Island Station, California State Fisheries Laboratory
for the quantity of preserved specimens from California and Lower California, and for
interesting notes on the ecology of several species. David and Nevada Schmidt collected
many lots on the west coast of Florida. Piet Kaas, the Hague, Holland and P. Rancurel,
Office de la Recherche Scientifique Outre-Mer, Paris, France, sent material from their
countries.

Among others who sent material I would like to thank the following: S. Stillman
Berry, Edward P. Baker, A. H. Cahn, Pedro de Mesa, Walter J. Eyerdam, Harry and

Western Atlantic JOHNSONIA, No. 33 19

Kathleen Johnstone, Roy Latham, Margaret Teare, Margaret Teskey, Gertrude Thorn-
ley and Gertrude Weber.

All photographs, unless otherwise stated, are the work of Frank White, Biological
Laboratories, Harvard University. The drawings were made by the author with the aid
of a camera lucida.

Subfamily PHoLADINAE
Genus Barnea Risso

Shells white, more or less elliptical in outline, and beaked or rounded anteriorly. Ac-
cessory plate consisting of a simple, calcareous, lanceolate protoplax. Umbonal reflec-
tions simple, closely appressed to or raised slightly above the surface of the umbos, but
free anterior to the umbos. Space below the umbonal reflections not septate. Pedal gape
variable, ranging from a narrow slit to a broad oval. Sculpture consisting of concentric
ridges and radial ribs which may cover the entire shell or may be reduced on the posterior
slope.

The genus Barnea Risso is known from the temperate and tropical seas throughout
the world.

Genotype, Barnea spinosa Risso (= Pholas candida Linné), monotypic.

There are two subgenera in the Genus Barnea and they may be differentiated as follows:

Shells rounded anteriorly Barnea s.s.
Shells beaked anteriorly Anchomasa

Subgenus Barnea Risso

Barnea ‘Leach’ Risso 1826, Histoire Naturelle de 1’Europe Méridionale 4, p. 376 (genotype, B. spinosa
Risso, monotypic).

Barnia Gray [sic] 1840, Synopsis Contents British Museum ed. 42, p. 150 [nomen nudum]; Gray 1842,
Synopsis Contents British Museum, ed. 44, p. 76 [defined but no species listed].

Barnia Leach [sic] 1852, Synopsis of the Mollusca of Great Britain, London, p. 254 (genotype, Barnea can-
dida Linné, monotypic).

Holopholas Fischer 1887, Manuel de Conchyliologie, Paris, p. 1133 (genotype, Barnea candida Linné, [here
designated | ).

Shells white, rounded at both ends, having a slit-like pedal gape and only a slight pos-
terior gape. Umbonal reflections simple and closely applied to the surface of the umbos.

No species of this subgenus is known from the Western Atlantic or the Eastern Paci-
fic. However, a description of the subgenus with a description and figures of Barnea
candida Linné are included for the sake of comparison. Other species in this subgenus
are found in West Africa and the Indo-Pacific.

Barnea (Barnea) candida Linné
Plates 7-8

Pholas candida Linné 1758, Systema Naturae, edition 10, p. 669; Hanley 1855, Ipsa Linnaei Conchylia,
Dp: 25:

Pholas papyraceus Spengler 1792, Skrivter Naturhistorie Selskabet 2, pt. 1, p. 92: non papyracea ‘Solander’
Turton 1822.

Barnea spinosa Risso 1826, Histoire Naturelle de 1’Europe Méridionale 4, p. 376.

Pholas dactyloides Delle Chiaje 1829, Memoire Animali senza Vertebrae Regno Napoli 4, p. 206, pl. 65,
figs. 9-12; non Lamarck 1818.

20 JOHNSONIA, No. 33 Barnea

Barnia [sic] candida Linné, Gray 1851, Annals and Magazine Natural History (2) 8, p. 382.

Barnea candida Linné, Bucquoy, Dautzenberg and Dollfus 1896, Mollusques Marins du Roussillon 2, p.
615, pl. 88, figs. 1-7.

Pholas candida var. subovata Jeffreys 1865, British Conchology 3, p. 108.

Pholas cylindrica J. Sowerby 1818, Mineral Conchology of Great Britain 2, p. 223, pl. 198 (Crag).

Pholas candida subovata cylindracea Marshall 1914, Journal of Conchology 14, no. 7, p. 207 (Torre Abbey,
Torquay, England).

Barnea candida cylindrica “Marshall? Lamy 1925, Journal de Conchyliologie 69, p. 40 [error for cylindracea
Marshall}.

Pholas costulata Goodall 1890, Transactions Norfolk and Norwich Naturalists Society 5, pt. 1, p. 79, text
fig. (Hill Head near Gosport, England) | We have not seen this publication]; Walsingham 1916, Proceedings
Malacological Society London 12, p. 61, text figure.

Distinctive characters. Shel usually 3 inches or less in length, rounded anteriorly, with
the sculpture extending over the entire shell and with a single dorsal plate, a protoplax.

Description. Shell subelliptical in outline, reaching about 68 mm. (about 2# inches) in
length and 25 mm. (about 1 inch) in height, thin, rounded anteriorly, tapering posteri-
orly and gaping slightly at both ends. Color a dull chalky white. Umbos prominent and
located near the anterior + of the shell. Umbonal reflections closely appressed over the
umbo but free anterior to the umbo. Sculpture consisting of concentric ridges and radial
ribs. Concentric ridges very strong on the anterior slope, becoming weaker over the disc
and on the posterior slope. Radial ribs rather weak, being expressed mainly by radial
arrangement of the rows of imbrications which are produced where the concentric ridges
and radial ribs cross. Imbrications particularly strong on the anterior slope but clearly
visible, though weak, the entire length of the shell. Sculpture below the umbonal re-
flection consisting of fine growth lines. Interior of the shell white and glazed. Surface
sculpture visible on the inner surface of the shell as faint furrows and pits. Pallial sinus
extending inward to about 4 the distance to the umbo. Muscle scars faint. Apophysis
small, short, slightly flattened and curved. Protoplax elongate oval, broadly arched, and
with a central groove extending its entire length. Posterior end of the protoplax sharply

Plate 7. Barnea candida Linné. Figs. 1-3. Bréhec, Cotes du Nord, France (all natural size). Fig. 1. Dorsal
view with protoplax in normal position. Fig. 2. Internal view showing apophysis, muscle scars and pallial
sinus. Fig. 3. External view of valve showing umbonal reflection which is free anterior to the umbos and
closely appressed over the umbos.

Western Atlantic JOHNSONIA, No. 33 21

bent downward, nucleus posterior and concentric growth lines faint. Periostracum thin,
straw-yellow and deciduous.

Siphons united, about equal in size and may be extended 4 to 5 times the length of
the shell. ‘They are pale brown in color and minutely papillose. he anterior portion of
the siphons with a darker brown periostracal sheath which is joined to and extends over
the posterior portion of the shell. Siphons tipped with a narrow band of red-brown. In-
current siphon with 10 to 12 large and several small cirri surrounding the opening. These
cirri continue internally as ridges which extend well down in the siphons. ‘The red-brown
coloring also extends internally largely between the ridges. Excurrent siphon lacking
cirri and with only a very narrow color band internally. Foot white, oblong-lanceolate
in outline and truncate. Pedal gape long and narrow, mantle white. The above descrip-
tion was made from preserved animals.

length height ratio h:1]

68.2mm. 25.0mm. 2.7 Cintra Bay, Rio de Oro, Africa

56.0 19.4 2.8 Ostende, Belgium

5280 19.0 a Castle Rocks, St. Andrews, Fife, Scotland

Types. According to Hanley (1855, Ipsa Linnaei Conchylia, p. 25) Linné’s specimens
of Pholas candida are still in the Linnean collection. The locality given by Linné was
Europe and America, his reference was to Lister 1678, Historiae Animalium Angliae,
p. 198, pl. 5, fig. 39. The type locality is here restricted to the Tees River, Middles-
brough, England, the locality given by Lister.

The locations of type specimens of most of the species now considered synonyms of
Barnea candida Linné are unknown.

Remarks. Barnea candida Linné, similar to all other species in this genus and family,
is rather variable in size, proportions and condition of sculpturing depending upon the
substratum in which it is boring. ‘This undoubtedly has been responsible for the large num-
ber of synonyms noted above. This species is known to bore into a variety of substrata
ranging from loose sand to rock.

Through the kindness of Professor C. M. Yonge and Dr. J. M. Dodd of the Univer-
sity of Glasgow, we have received preserved specimens from the Gatty Marine Labora-
tory, St. Andrews, Scotland. At this station Barnea candida Linné bore into mudstone,
a form of shale which is quite plastic when wet. They live in the same area as Zirfaea
crispata Linné, but at a higher tide level and their burrows generally are horizontal while
those of Zirfaea are nearly always vertical.

Deshayes (1846, Exploration Scientifique de |’ Algerie, Histoire Naturelle des Mollus-
ques, Liv. 6, pl. 9) has figured beautifully the general morphology of the soft parts of
this species, and Bucquoy, Dautzenberg and Dollfus give a very complete bibliography
prior to 1896.

Range. From northern Norway south to the southern coast to France, the eastern
Mediterranean, and on the African coast south to Cap Blane, Sénégal (M. Nicklés, 1950).

Specimens examined. Norway: Malvik, Trondheim Fjord; Beian, Orlandet, 'Trond-
heim (both MCZ). Scornanp: Oban (MCZ; USNM). EnNGianp: Huntstonton

22 JOHNSONIA, No. 33 Barnea

(USNM); Scarborough: Kent; Eastbourne, Sussex ; Teignmouth; Cornwall (all MCZ);
Poole (USNM). NerHErRLANDs: Scheveningen (MCZ). BELcium: Nieuport (W. J.
Eyerdam); Ostende (MCZ). France: Bréhec, Cotes du Nord; Nantes; Cette (all
MCZ); Gulf of Gascogne (USNM). Arrica: Cintra Bay, Rio de Oro (MCZ).

iii
AM
ING

CE

~~ ~—S—>

K«

we

Plate 8. Protoplax of Barnea. Figs. 1-3. Barnea subtruncata Sowerby, Liddo Isle, Newport Bay, California.
Fig. 1. Dorsal view showing the tapering sides, the concentric growth lines and the nucleus which is lo-
cated well in from the posterior margin. Fig. 2. Ventral view. Fig. 3. Side view to show the curvature.
Figs. 4-6. Barnea candida Linné, Bréhec, France, Fig. 4. Dorsal view showing the posterior nucleus and
the distinct median groove. Fig. 5. Ventral view showing the groove expressed internally as a ridge. Fig.
6. Side view. Figs. 7-9. Barnea truncata Say, Third Cliff, Scituate, Massachusetts. Fig. 7. Dorsal view
showing the nearly parallel sides, the concentric growth lines and the nucleus which is located close to the
posterior margin. Fig. 8. Ventral view. Fig. 9. Side view. Figs. 10-12. Barnea parva Pennant, Plymouth,
England. Fig. 10. Dorsal view showing the growth lines and the location of the nucleus. Fig. 11. Ventral
view. Fig. 12. Side view. Figs. 13-15. Barnea lamellosa d’Orbigny, Puerto Militar, Bahia Blanca, Buenos
Aires, Argentina. Fig. 13. Dorsal view showing the weak concentric growth lines, the posterior nucleus
and the posterior lobes. Fig. 14. Ventral view showing the longitudinal ridge. Fig. 15. Side view showing
the curvature and the upturned lateral margins.

Subgenus Anchomasa Leach
Anchomasa Leach 1852, A Synopsis of the Mollusea of Great Britain, London, p. 253.

Shell beaked anteriorly, with a large oval pedal gape extending back at least as far as
the umbo, rounded to truncate posteriorly with a moderate to large and variable posterior

Western Atlantic JOHNSONIA, No. 33 23

gape. Umbonal reflection simple and usually closely applied to the surface of the umbo,
free anteriorly.

Subgenotype, Anchomasa pennantiana Leach (= Pholas parva Pennant), monotypic.

Barnea (Anchomasa) parva Pennant
Plates 8-9

---

Pholas parva Pennant 1777, British Zoology 4, p. 77, pl. 40, fig. 13 (English shores); non parva Sowerby
1834; non parvus Donovan 1800.

Pholas crenulatus “Solander’ Spengler 1792, Skrivter Naturhistorie Selskabet 2, pt. 1, p. 98.

Pholas dactyloides Lamarck 1818, Histoire Naturelle des Animaux Sans Vertébres 5, p. 445 (British Seas).

Pholas tuberculata Turton 1822, Conchylia Insularum Britannicarum, p. 5, pl. 1, figs. 7-8 (Torbay, England).

Pholas ligamentina Deshayes 1839, Traité Elémentaire de Conchyliologie 1, pt. 2, p. 80, pl. 3, figs. 11-12
(Seas of Europe).

Barnia [sic] parva Pennant, Gray 1851, Annals and Magazine of Natural History (2) 8, p. 382.

Anchomasa pennantiana Leach 1852, Synopsis of the Mollusca of Great Britain, p. 253 (Kingsbridge Estuary
and Sussex, England).

Zirfaea callosa ‘Lamarck’ Chenu 1862, Manuel de Conchyliologie 2, p. 6, figs. 24-25.

Barnea ( Anchomasa) parva Pennant, Tryon 1862, Proceedings Academy Natural Sciences Philadelphia, p.209.

Barnea parva quadrangula Jeffreys 1865, British Conchology 3, p. 110 (England).

Holopholas (Anchomasa) parva Pennant, Fischer 1886, Manuel de Conchyliologie, p. 1133.

Pholas duboisi Locard 1892, Les Coquilles Marines des Cétes de France, p. 246 (coast of France and Eng-
lish Channel).

Pholas (Barnea) parva var. major Pallary 1900, Journal de Conchyliologie 48, p. 413, fig. 19 (St. Thérése,
Oran [Algeria] ).

Distinctive characters. Shell strongly beaked anteriorly, narrowly rounded posteriorly,
solid, with a large rounded condyle and a lanceolate protoplax.

Description. Shell reaching 38 mm. (about 14 inches) in length and about 16 mm.
(about $ inch) in height, rather strong in structure, beaked anteriorly, rounded posteri-
orly, gaping at both ends and with the sculpture disappearing on the posterior slope.
Color a dull chalky white. Umbos prominent and located near the anterior third of the
shell. Umbonal reflections narrow, rather thick and raised above the umbos. Sculpture
consisting of radial ribs and concentric ridges. Concentric ridges strongly laminated an-
teriorly, becoming weak on the disc and disappearing on the posterior slope. Radial ribs
closely set on the anterior slope, barely visible on the disc and lacking on the posterior
slope. Imbrications are produced where the concentric ridges and radial ribs cross one
another. In most specimens the radial arrangement of the imbrications is the only evi-
dence of the ribs. Sculpture beneath the umbonal reflection consisting only of laminated

Plate 9. Barnea parva Pennant. Fig. 1. Nantes, France (2x). Showing the large knob-like condyle. Fig.
2. Tenby, Pembroke, Wales (2x).

24 JOHNSONIA, No. 33 Barnea

ridges. Interior of shell white and glazed. Pallial sinus wide and deep, extending in-
wardly almost to the umbo. Pallial line and muscle scars barely visible on the specimens
examined. Radial ribs visible internally on the anterior slope. Condyle knob-like, large
and heavy and extending inward so that when the two valves are in juxtaposition they
are quite widely separated along the dorsal margin. Apophysis narrow and thin. Proto-
plax lanceolate. Periostracum thin, and straw-yellow in color.

Siphons united and may be extended 1 to 2 times the length of the shell. They are
thickly covered with small papillae, fawn-brown in color and have a narrow whitish
band surrounding the openings. Incurrent siphon nearly three times the diameter of the
excurrent siphon and with three large and several small ridges extending well down
within. Excurrent siphon smooth internally. Internal surface of both siphons white to
pale yellow. Foot and mantle white to pale yellow. Foot large, nearly circular in out-
line and truncate. The above description of the soft parts is based on preserved speci-
mens received through the kindness of Dr. F. S. Russell, Director of the Plymouth
Laboratory, England.

length height ratio h:]
38.0 mm. 16.0 mm. 2.4 Nantes, France
S5n5 16.0 Dee Tenby, Pembroke, Wales
25D 13.5 1.8 Plymouth, England
24.0 12.0 2.0 Plymouth, England
DONE 1250 1.8 Tenby, Pembroke, Wales

Types. The type of Barnea parva Pennant is probably in the British Museum. Ac-
cording to Sherborn 1940, the collections of Pennant were deposited in the British Mu-
seum in 1912. The type locality is Abergelli [Abergele] in Denbighshire, Wales. The
locations of the type specimens of most of the species now considered synonyms of Barnea
parva Pennant are unknown.

Remarks. Barnea parva Pennant is the type of the subgenus Anchomasa and for this
reason we include a description of it here. It is a rather small and variable species with
an unusually heavy shell for a Barnea of this size. It is characterized by its pronounced
beak, its closely packed concentric ridges and radial ribs, its prominent dorsal condyles
and the narrowly rounded posterior margin. Barnea parva Pennant most nearly approx-
imates Barnea lamellosa d’Orbigny of Argentina. The posterior margins of both species
are rounded and tapering and in both, the hinge area is somewhat knob-like. PBarnea
parva Pennant, however, has a much heavier, more solid shell, with close-set laminated
ridges and a simple non-lobed protoplax. From Barnea truncata Say which has also been
found in the Eastern Atlantic on the African coast, B. parva Pennant is distinguished
by its rounded posterior margin, more pronounced beaks, heavy shell, and more knob-
like development of the hinge area.

Barnea parva Pennant is apparently a rather rare species with a somewhat restricted
range. Forbes and Hanley 18538 state that while found with Barnea candida, Pholas
dactylus and Zirfaea crispata, it is always far less abundant. It bores into soft sandstone,
stiff clay and decaying waterlogged wood, the general shape and size of the specimen
depending upon the hardness of the material into which it is boring. It is this variation
that has, no doubt, been responsible for the large number of synonyms noted above. The

Western Atlantic JOHNSONIA, No. 33 25

specimens which Pennant cites in his original description as coming from wood taken at
Pensacola, Florida, were no doubt malformed specimens of B. truncata. When boring
in such hard material they often look very much like parva, but never are as heavy in
structure nor do they possess the large knob-like condyles.

Dr. M. V. Labour’ states that this species is common in the rocks at Rum Bay, Ply-
mouth, England and that they appear to breed in summer and early autumn. She reared
the larvae to the first shell stage and described and figured them. Like most bivalves, the
larval shell is very small, being only .08 mm. in length and extremely simple in struc-
ture. When .32 mm. in length, it begins to assume the adult form, developing an an-
terior portion which is imbricated and suitable for boring, and a posterior portion for the
protection of the siphons.

Range. Southern British Isles south to Oran, Algeria (Pallary).

Specimens examined. WALES: Tenby, Pembroke (MCZ). Excianp: Tor Bay, Devon
(Charleston Museum); Brighton, Sussex (W.J. Eyerdam); Plymouth (Plymouth Lab. ).

France: Nantes (MCZ). La Rochelle (USNM).

Barnea (Anchomasa) lamellosa d’ Orbigny
Plates 8 and 10

Pholas lamellosa d°Orbigny 1846, Voyage Amérique Méridionale 5, pt. 3, Mollusques, p. 498, pl. 77, fi
20-21 (shore of Patagonia to the south of Rio Negro).

Barnea (Anchomasa) subtruncata var. lamellosa d’Orbigny, Lamy 1925, Journal de Conchyliologie 69, p. 82.

Barnea (Anchomasa) subtruncata lamellosa d’Orbigny, Carcelles 1944, Revista Museo de la Plata (n.s.) Zool-
ogia 3, p. 295, pl. 14, figs. 108-109; Carcelles 1950, Anales del Museo Nahuel Huapi 2, p. 82.

gs.

Distinctive characters. Shell white, beaked anteriorly, rounded posteriorly, with the
umbonal reflections free for their entire length, with a large rounded condyle and a single
dorsal plate, the protoplax, which is strongly keeled and eared posteriorly.

Description. Shell reaching about 43 mm. (1# inches) in length and about 184 mm.
(about # inch) in height, thin, frail, pointed anteriorly, rounded posteriorly, gaping at
both ends, and with the sculpture disappearing on the posterior slope. Color a dull chalky-
white. Umbos prominent, partially covered by the umbonal reflections and located near
the anterior third of the shell. Umbonal reflection free for its entire length, the space

Plate 10. Barnea lamellosa d’Orbigny, Mar del Plata, Argentina (2x). Fig. 1. Showing the free umbonal re-
flection and high posterior slope. Fig. 2. Showing the muscle scars and apophysis.

GS

‘Labour, M.V. 1933, Journal of the Marine Biological Association of the United Kingdom 23, no. 1, pp.
132-130.

26 JOHNSONIA, No. 33 Barnea

between it and the umbo being very small in young specimens and increasing slightly in
older shells. Sculpture consisting of rather strong concentric ridges and moderate to
rather weak radial ribs. Concentric ridges laminated anteriorly, becoming smoother and
weaker posteriorly until they are nearly obsolete on the posterior slope. Radial ribs gen-
erally rather weak and in adult specimens usually completely lacking on the posterior
slope. Imbrications are produced where the radial ribs and concentric ridges cross. The
strength of these imbrications varies greatly depending on the age of the specimen and
the strength of the ribs. On young specimens (up to 12 mm. in length) they may even
be present on the posterior slope. In the rather small series available for study the radial
arrangement of the imbrications was the only evidence of the existence of the radial
ridges, there being no trace of them between the crests of the concentric ridges. Sculp-
ture below the umbonal reflection composed of crowded laminated growth lines. Interior
of shell white and glazed, with the concentric lines and ridges showing through the shell.
Pallial sinus nearly as wide as the shell is high and extending inward nearly half the dis-
tance to the umbo. Apophyses narrow, flattened, and extending downward about } of
the way to the ventral edge of the shell.

Protoplax lanceolate with a posterior nucleus and well marked growth lines, pointed
anteriorly, broad and eared posteriorly, strongly keeled, and with a strong hook at the
posterior end which fits over the prominent umbos. Pedal gape rounded anteriorly,
pointed posteriorly and extending backward about one half the length of the shell. Peri-
ostracum very thin, straw-yellow and deciduous.

The siphons are united and may be extended 4 to 5 times the length of the shell. The
covering of the siphonal tubes in preserved material is medium to dark-brown in color
and papillose. There are 8 to 10 short stout papillae surrounding the end of the incurrent
siphon. These papillae extend inside as marked white ridges almost the entire length of
the incurrent siphon, the spaces between the ridges a dark red-brown. The excurrent
siphon lacks papillae and is smooth within. The strongly muscular mantle collar surround-
ing the foot and forming the pedal gape is notched anteriorly at the ante-umbonal gape.

length height ratio h:]
42 mm. 19mm. De Mar del Plata, Argentina
38 15 DG. Puerto Quequen, Buenos Aires, Argentina

Types. The holotype of Barnea lamellosa d’Orbigny is in the British Museum, accord-
ing to Gray 1854, List of the Shells of South America in the Collection of the British
Museum Collected and Described by M. Alcide d’Orbigny, p. 55. The type locality is
the mouth of the Rio Negro, Argentina.

Remarks. Barnea lamellosa d’ Orbigny is close in its relationship to B. truncata Say.
It differs, however, in being more rounded rather than truncate posteriorly, in having the
umbonal reflection free for its entire length and in having the protoplax strongly keeled,
more pointed anteriorly and eared posteriorly. Barnea lamellosa d’Orbigny also appears
to be a much smaller species and to have much weaker sculpture with more closely set
concentric ridges than B. truncata Say. See also remarks under B. parva Pennant.

Though specimens of B. /amellosa are exceedingly rare in museum collections in this
country, it is probably not a rare species. It has a limited distribution and is further re-
stricted by its specialized habitat of mud and peat areas. Through the kindness of A.
Carcelles we have had specimens from several localities in Argentina for study. We agree

Western Atlantic JOHNSONIA, No. 33 Q7

~t

with Carcelles that this species is not the same as the one found in Peru, as has been
stated by several authors. This species has a characteristic eared protoplax, a heavy con-
dyle and a much more rounded posterior margin than B. subtruncata Sowerby, the East-
ern Pacific species.

Range. From off Cabo Polonio, Uruguay south to Golfo Nuevo, Argentina.

Specimens examined. Urucuay: Off Cabo Polonio (S. Lat. 84°42’: W. Long. 54°10’)
(A. Carcelles). ARGENTINA: Mar del Plata; Bahia Blanca; Puerto Quequen; Canal la
Manuelita; Puerto Belgrano; Bahia San Blas, all in the State of Buenos Aires; 5 miles
south of Punta Medano, Rio Negro; Puerto Madryn and Golfo Nuevo, Chubut (all

A.Carcelles). Rio Negro (USNM).

Barnea (Anchomasa) truncata Say
Plates 8 and 11 to 13
Pholas truncata Say 1822, Journal Academy Natural Sciences Philadelphia 2, p. 321 (southern coast of
United States).

Pholas (Cyrtopleura) truncata Say, Tryon 1862, Proceedings Academy of Natural Sciences Philadelphia 14,
p. 202.

Barnea truncata Say, Dall 1898, Transactions Wagner Free Institute of Science, Philadelphia 3, part 4, p.816.
Barnea (Cyrtopleura) truncata Say, Lamy 1925, Journal de Conchyliologie 69, p. 87.

Distinctive characters. Shell beaked anteriorly, truncate posteriorly, sculpture greatly
reduced or entirely lacking on the posterior slope, and with a single dorsal plate, the
protoplax.

Plate 11. Barnea truncata Say. Woods Hole, Massachusetts (natural size). Living specimens photographed in
test-tubes. Fig. 1. Ventral view of specimen ina more or less relaxed condition with the longitudinal mus-
cles slightly contracted and showing the extent to which the valves can be separated. Fig. 2. With foot
well extended and the longitudinal and circular muscles of the siphons moderately contracted. Fig. 3.
Dorsal view showing the protoplax in place and the extent to which the valves can be separated dorsally.
Fig. 4. Side view showing the extended foot and the circular muscles of the siphons contracted.

28 JOHNSONIA, No. 33 Barnea

Description. Shell reaching 70 mm. (about 24 inches) in length and 25 mm. (about 1
inch) in height, light in structure, beaked anteriorly, gaping at both ends and with the
sculpture disappearing on the posterior slope. Color a dull chalky-white. Umbos prom-
inent and located near the anterior third of the shell. Umbonal reflections rather closely
appressed over the umbos but free anteriorly. Sculpture consisting of radial ribs and con-
centric ridges. The concentric ridges strong and laminated anteriorly, becoming gradually
weaker posteriorly until they are nearly obsolete on the posterior slope. Radial ribs fairly
strong on the anterior slope, diminishing in strength on the disc and completely lacking
on the posterior slope. Imbrications are produced where the radial ribs and concentric
ridges cross. In some specimens, the radial arrangement of these imbrications is the
only evidence of the existence of the radial ribs. The degree of sculpturing below the
umbonal reflection varies greatly, even among specimens from the same locality, ranging
from very weak growth lines to sculpture almost as strong as that on the anterior slope.
Interior of the shell white and glazed. Pallial sinus nearly as wide as the shell is high and
extending within nearly half the distance to the umbo. Pallial line and muscle scars
clearly indicated especially on older specimens. Apophyses long, narrow, curved and
blade-like, occasionally broadening slightly at the free end. Protoplax broadly lanceo-
late, with a posterior nucleus and well marked growth lines. It is curved downward at its
posterior extremity to fit over the umbo. Periostracum thin, straw-yellow to light-brown
in color and deciduous. .

Siphons united and enclosed in a tough, brown, papillose sheath, the incurrent siphon
being a little larger than the excurrent. They may be extended 10 to 12 times the length
of the shell, and, when fully extended, the periostracal covering is so stretched that they
appear a light brownish-gray to white. However, on contraction they become a uniform
dark brown their entire length. Incurrent siphon fringed with 8 to 10 short, stout, un-
branched papillae. Between these papillae there are brown to mahogany-red markings
which extend down inside the siphon. The excurrent siphon lacks the papillae but usu-
ally possesses the brown internal markings. The dark periostracal covering extends an-
teriorly between the valves on the ventral surface so that only the immediate area sur-
rounding the foot is white. Foot white, oval in outline and truncate.

length ’ height ratio h:]
70.0 mm. 27.0 mm. 2G Duxbury, Massachusetts
66.5 26.5 oD Sarasota, Florida
62.0 2O7D Dev Mattituck, Long Island, New York
59.0 25.0 2.4 Duxbury, Massachusetts
54.0 23.0 2.3 Coty pe
47.0 as: 259 Marion, Massachusetts
44.5 20.0 2.8 Newport, Rhode Island

Types. A probable cotype of Pholas truncata Say is in the Academy of Natural
Sciences Philadelphia, no. 50775, a specimen received from Mrs. Say. Say’s type locality
was simply the southern coast of the United States. We here restrict the type locality to

Charleston, South Carolina, a region from which Say received much material.

Remarks. Barnea truncata Say is very close in its relationship to B. subtruncata Sowerby
from the Eastern Pacific. It is, however, generally a smaller, more fragile shell, with less
inflated valves and shorter post-umbonal length. In addition, the protoplax is less taper-
ing. In most of the above shell characters, however, there are intergrades, though ina

Western Atlantic JOHNSONIA, No. 33 29

large series the two forms appear quite distinct. For additional remarks on relationship
to the west coast form see under B. subtruncata Sowerby. See also remarks under Barnea
lamellosa d’Orbigny.

Specimens of Barnea truncata Say are found in mud, clay and peat deposits along the
coast from Maine to Brasil wherever strictly marine conditions exist. The distribution
of this species north of Cape Cod is extremely local and specimens from Maine, the
northern limit of its range, are exceedingly rare. The species again becomes rare on the
southern end of its range. It would appear that in this area they are probably living in
deeper water, and being deep borers the shells seldom get washed ashore. Harold W.
Harry’ reports that Barnea truncata was found commonly in the beach drift at Timbalier
Island and Caminada, Louisiana. However, to our knowledge, no living specimens have
been collected in the northern Gulf or in the West Indies.

Large specimens of B. truncata may bore holes nearly a foot or more in depth and
though usually found inhabiting the intertidal region they occasionally may be found at
considerable depths. Small specimens of this species were found in a piece of waterlogged
wood tossed up on the beach at Cape Canaveral, Florida. These specimens were stunted
and malformed, the wood being much harder than the material into which they normally
bore. A similar malformed specimen in the collection of the Charleston Museum was
labeled as boring in rock in Charleston Harbor. This was probably the hard marl or phos-
phate rock of that area, a substance much too hard for B. truncata to bore into success-
fully. The beaks of these specimens were greatly produced, the rows of imbrications ex-
ceedingly close together, and the posterior slope much reduced in size. Specimens from
several localities on the west coast of Florida in the vicinity of Sarasota appear to be some-

Plate 12. Barnea truncata Say. Figs. 1, 3. Newport, Rhode Island. Fig. 2. Duxbury, Massachusetts. Selected
series to show variation in the length-height proportion (all natural size).

' Harold W. Harry 1942, Occasional Papers Marine Laboratory, Louisiana State University, Baton Rouge,
Louisiana, no. 1, p. 3.

30 JOHNSONIA, No. 33 Barnea

what longer in proportion to their height and to have the posterior margin more rounded
than the typical form. This may be due to ideal conditions of the substrata and a good
food supply so that the specimens grew more rapidly than usual.

Edward Sylvester Morse found that the animals when cooked were quite edible, but
had a peculiar smoky taste.

Addison E. Verrill (1873), appears to have confused the soft parts of Barnea truncata
and Zirfaea crispata, for he describes the siphons of truncata as *‘generally yellowish-
white except at the very end, where they are blackish or brownish; the orifices and pap-
illae are also variously marked with purplish brown or dark brown. The dark coloration
at the end of the siphon tubes is doubtless for purposes of protection from predaceous
fishes, crabs, ete.” This description fits perfectly the siphons of Z. crispata, but the
siphons of B. truncata are a uniform grayish-brown their entire length.

Range. WerstERN ATLANTIC: Maine (C. W. Johnson) south to the Gulf of Mexico,
probably in restricted localities throughout the West Indies, and south to Sepetiba Bay,
Rio de Janeiro, Brasil.

Eastern AruLantic: From Bel Tir, Dakar, Sénégal south to Accra, Gold Coast.

Specimens examined. WESTERN ATLANTIC: Massachusetts: Salem (MCZ; USNM);
Chelsea Beach (USNM); Scituate; Duxbury; Marion; Dennisport; West Harwich;
Hyannis: Woods Hole (all MCZ); Lagoon Pond, Marthas Vineyard; Cuttyhunk Island
(both G. Moore); New Bedford (ANSP). RuopE IsLanp: Newport; Warren (both

Plate 13. Barnea truncata Say. Figs. 1-2. Cotype, from South Carolina (about 1.2x). Fig. 1. Showing mus-
cle scars and pallial sinus. (Apophysis lost.) Fig. 3. Accra, Gold Coast, Africa (natural size).

Western Atlantic JOHNSONIA, No. 33 31

MCZ): Bristol (AMNH). Connecricur: New Haven (USNM); Branford (MCZ).
New York: Orient and Barren Island, Long Island (both R. Latham); Mattituck
Creek, Mattituck and Plum Island, Long Island (both AMNH); Lido Beach, near
Atlantic Beach, Long Island (M. K. Jacobson); Staten Island (MCZ: USNM); Univ.
of Michigan). New Jersey: Point Pleasant; Sea Isle City (both ANSP); Atlantic
City (USNM; ANSP); Cape May (MCZ; USNM; ANSP; Univ. of Michigan):
Great Egg Bay (USNM). Vireinia: Hog Island, Eastern Shore (USNM). Norru
Caro.ina: Fort Macon (MCZ); Beaufort (MCZ; USNM). Sourn Carotina: Myrtle
Beach; Pawleys Island; Isle of Palms; Sullivans Island; Folly Island; Magnolia Beach
(all Charleston Museum); Charleston Harbor (Charleston Museum; USNM). Georeia:
Sea Island, St. Simons Island (USNM; ANSP); Savannah (USNM). FLoripa: St.
Augustine (USNM); Daytona Beach; Cocoa (both MCZ); Flagler Beach; Cape Sable
(both Univ. of Miami); Pavilion Key; Lossmans Key (both D. and N. Schmidt); San-
ibel Island (MCZ; USNM); Sarasota; Anna Maria Island (both R. Stewart); Braden-
ton Beach (A. Koto; D. and N. Schmidt); Boca Ciega Bay, St. Petersburg (Univ. of
Michigan); Clearwater Beach (ANSP); Port St. Joe (A. Merrill). Texas: Sand Point,
Calhoun Co.; Carancahua Bay, Matagorda Bay (both USNM). Hispaniota: Monte
Cristi, Santo Domingo (MCZ). TrRintbap: Moruga(H. G. Kugler). Brasi_t: Parada
Sahy, Sepetiba Bay, Rio de Janeiro (H. S. Lopes).

EasrerN ATLANTIC. SENEGAL: Bel Tir, Dakar (Institut Franeais d’ Afrique Noire).
Gop Coast: Accra (MCZ).

Barnea (Anchomasa) subtruncata Sowerby
Plates 8 and 14 to 16

Pholas subtruncata Sowerby 1834, Proceedings Zoological Society London, p. 69 (Insula Platae, Colum-
biae Occidentalis [ Ecuador].

Pholas spathulata Deshayes 1843, Magasin de Zoologie (par Guérin-Méneville) (2) 5, Mollusques, pl. 79
(Seas of Chile); non Pholas spathulata Sowerby 1849.

Barnea pacifica Stearns 1871, Conchological Memoranda no. 7, p. 1 (Preliminary description); Stearns 1873,
Proceedings California Academy of Science 5, p. 81, pl. 1, fig. 6, 6a—c (Alameda, San Francisco Bay, Cali-
fornia); Stearns 1891, Proc. United States National Museum 14, p. 314.

Barnea (Cyrtopleura) spathulata Deshayes, Lamy 1925, Journal de Conchyliologie 69, p. 89.

Barnea pacifica Stearns, Hertlein and Strong 1950, Zoologica 35, p. 248.

Distinctive characters. Shell beaked anteriorly, broadly rounded to truncate posteriorly,
with the sculpture usually lacking on the posterior slope, and with a long narrow acumi-
nate protoplax.

Description. Shell reaching 67 mm. (22 inches) in length and 27 mm. (about 1 inch)
in height, light in structure, beaked anteriorly, gaping at both ends and with the sculp-
ture disappearing on the posterior slope. Umbos prominent, usually located near the
anterior fourth of the shell. Umbonal reflections closely appressed over the umbos, free
anteriorly. Sculpture consisting of concentric ridges and radial ribs. Concentric ridges
low, usually rather widely spaced, laminated and scalloped anteriorly, becoming much
weaker on the disc and evident only as growth lines on the posterior slope. Radial ribs
moderate in strength on the anterior slope, weak on the disc and completely lacking on
the posterior slope. Broad and rather weak imbrications are produced where the concen-

$2 JOHNSONIA, No. 33 Barnea

tric ridges and radial ribs cross one another. Sculpture beneath the umbonal reflection
generally consists of crowded and weak concentric ridges. Interior of shell white and
glazed. Muscle scars and pallial lines clearly indicated. Pallial sinus broad and extend-
ing inward nearly half the distance to the umbo. Apophyses long, narrow and curved
inwardly. Protoplax lanceolate in outline, usually rather broad and truncate posteriorly,
acuminate anteriorly, with a posterior nucleus and well marked growth lines. Periostra-
cum thin on the anterior slope, rather heavier posteriorly and ranging from a straw-yellow
to a dark-brown in color.

Siphons enclosed in a sheath which ranges from a dark red-brown to dark-gray in color,
and usually with a narrow white tip at the posterior extremity, minutely papillose the
entire length. In some specimens the dark papillose covering is worn off on the tops of
the ridges, giving the specimen a mottled appearance. Incurrent siphon much larger than
the excurrent and with two large dorsal ridges and ten small ridges running the entire
length of the siphon internally. The interior of the siphon a deep mahogany-red posteri-
orly, becoming less intense as it extends forward. The exhalant siphon has one large
ventral ridge and eleven small ridges internally; the dark-mahogany coloring stopping
abruptly just a short distance inside the siphonal opening. These ridges terminate at the
opening of the siphon in unbranched papillae. The dark periostracum extends anteriorly
between the valves so that only the immediate area surrounding the foot is devoid of
periostracum and is white in color. he above description of the animal was based on
specimens preserved in alcohol.

length height ratio h:]
73.0mm. 21.0mm. 3.5 Alamitos Bay, California
68.0 25.0 Qi Anaheim Bay, California
66.5 25.0 2.6 as = ri
63.0 26.5 2.3 Lectotype of B. pacifica Stearns
62.0 31.5 2.0 Lido Island, Newport Bay, California
61.0 29°5 Qt Alamitos Bay, California
5acD 26.5 aa Anaheim Bay, California
5Deo 9555 ae Payta, Peru
Boo 25.0 2.1 Anaheim Bay, California

Plate 14. Barnea subtruncata Sowerby. Alamitos Bay, California (about natural size).
Photograph supplied by John E. Fitch, California Fisheries Laboratory.

Western Atlantic JOHNSONIA, No. 33 33

Types. The type of Pholas subtruncata Sowerby according to Mr. G. L. Wilkins of the
British Museum is no longer in existence. The type locality is Insula Platae, Colum-
biae Occidentalis [Isla la Plata, Ecuador]. The type of Pholas spathulata Deshayes is in
the Paris Museum according to Lamy (1925, p. 89) and in accordance with Lamy we
here restrict the type locality to Payta, Peru. The lectotype of Barnea pacifica Stearns
from Alameda, San Francisco Bay, California is in the United States National Museum
no. 74717, paratypes are in the California Academy of Science and the Museum of Com-
parative Zodlogy.

Plate 15. Barnea subtruncata Sowerby. Fig. 1. Lectotype of Barnea pacifica Stearns (= Barnea subtruncata
Sowerby) San Francisco Bay, Alameda, California (about 14x). Figs. 2-3. Copies of the original figures
of Barnea spathulata Deshayes from the Magazine de Zoologie (2) 5, p. 79 (= Barnea subtruncata Sower-
by), from Payta, Peru.

Remarks. Barnea subtruncata Sowerby is in all respects very closely related to B.
truncata Say of the Western Atlantic. Adult shells of B. subtruncata Sowerby are usu-
ally larger in size and are generally slightly longer posterior to the umbo. The protoplax
is usually narrower in proportion to its length and more acuminate anteriorly. In addi-
tion, the siphons of the west coast species are somewhat darker in color, as well as being
more strongly ridged and colored internally. ‘The siphons of B. truncata are nearly
smooth internally and usually have medium brown color markings.

Barnea subtruncata Sowerby appears to be the earliest name for this species. Unfor-
tunately the type is not in existence and Sowerby’s description is poor. He compares
B. subtruncata with B. parva of England, a species close in its relationship with the
present form, and so faras is now known there is only one species in the Eastern Pacific
that could be compared with the European species. ‘There seems to be no question that
B. spathulata Deshayes and B. pacifica Stearns refer to the same species, for both Stearns
(1891) and Dall (1909) refer to the Peruvian species as B. pacifica and give the range as
from California to Chile. Consequently on the basis of the material now available and
considering our present knowledge of the distribution of the group, these three names all
seem to refer to the same species.

Barnea subtruncata, like its Western Atlantic relative B. truncata Say, is an ex-
tremely variable species both in appearance and habit. These clams are found in mud,

34. JOHNSONIA, No. 33 Barnea

clay and peat deposits, but occasionally specimens bore into soft rock or waterlogged
wood. These specimens are always more or less stunted, with more produced beaks, more
closely set imbricated ridges and with the posterior slope reduced and more rounded. It
was probably such a specimen upon which Sowerby based his description, as he said it
was boring in soft stone.

Range. From Newport, Oregon south to Atacama Province, Chile (Gigoux 1934,
Revista Chilena de Historia Natural 38, p. 285).

Specimens examined. OREGON: Newport; Yaquina Bay (both ANSP). Catirornia:
Alameda, San Francisco Bay (USNM); San Francisco Bay (MCZ; ANSP); Anaheim
Landing (MCZ; USNM; ANSP); San Pedro Bay (MCZ; E.P.Chace); Lido Island,
Newport Bay (E. Baker); Alamitos Bay (J. E. Fitch; E. P. Chace); Bolinas (ANSP);
La Playa, San Diego (USNM). Mexico: Kino Bay, Sonora (ANSP); Mendina, Sin-
aloa (USNM). Gatapacos IsLanps: (USNM). Peru: Payta (ANSP).

Plate 16. Barnea subtruncata Sowerby. Fig. 1. Payta, Peru. Figs. 2-4. Anaheim Bay, California. Figs. 1-3.
These figures show the variation in the truncation of the posterior margin and the length of the post-
umbonal area. Fig. 4. Internal view to show the apophysis, muscle sears and the pallial sinus (all natural
size).

Genus Cyrtopleura T7yon
Cyrtopleura Tryon 1862, Proceedings Academy Natural Sciences Philadelphia, p. 201.

Shells white, more or less elliptical in outline and rounded or beaked anteriorly. Pedal
gape variable, ranging from a narrow slit to a broad oval. Accessory plates consisting of
a protoplax and a mesoplax. Protoplax entirely chitinous or only slightly impregnated
with calcium. Mesoplax transverse, in one or two pieces, calcareous and solid. Umbonal
reflections simple, rather narrow, raised well above the umbos and with posterior sup-
ports. Sockets are formed in the posterior supports for the reception of the forward ex-
tension of the mesoplax. Space below the umbonal reflection not septate.

Genotype, Pholas crucifera Sowerby (= Pholas cruciger Sowerby), subsequent desig-
nation, Stoliczka 1870.

Western Atlantic JOHNSONIA, No. 33 35

There are two subgenera in the genus Cyrtopleura and they may be differentiated as
follows:
Shell rounded anteriorly with a slit-like pedal gape Scobinopholas
Shell beaked anteriorly with an oval pedal gape Cyrtopleura s.s.

Subgenus Scobinopholas Grant and Gale

Pholas Lamarck 1801, Systeme des Animaux sans Vertébres, Paris, p. 127 (genotype, P. costata Linné,
monotypic); non Lamarck 1799; non Linné 1758.

Pholas “Linné’ H. and A. Adams 1856, The Genera of Recent Mollusca 2, p. 325 (genotype, P. costata
Linné, subsequent designation, Bayle 1880); non Pholas Linné 1758; non Lamarck 1799.

Scobina Bayle 1880, Journal de Conchyliologie 28, p. 242 (genotype, Pholas costatus Linné, original desig-
nation). [New name for Pholas H. and A. Adams; non Linné 1758. |

Scobinopholas Grant and Gale 1931, Memoirs San Diego Society Natural History 1, p. 431 [new name for
Scobina Bayle 1880; non Scobina Lepeletier 1825; non Wade 1917].

Subgenotype, Pholas costatus Linné, Bayle, original designation.

Shell rounded at both ends and having a long, narrow, slit-like pedal gape. Accessory
plates consisting of a triangular to T-shaped, thin, largely chitinous protoplax and a
heavy transverse calcareous mesoplax, which may be in one or two pieces. Umbonal re-
flections well separated from the surface of the umbos and supported at the posterior
margins where they bend downward and form sockets for the reception of the forward
extension of the mesoplax. Apophyses large, broad, and more or less spoon-shaped.

Cyrtopleura (Scobinopholas) costata Linné
Plates 17 and 18

Pholas costatus Linné 1758, Systema Naturae, ed. 10, 1, p. 669 (Europe).

Capulus shreevei Conrad 1869, American Journal of Conchology 5, p. 105, pl. 13, fig. 3 (Long Island, South
Carolina). [This is the apophysis only. |

Scobina costata Linné, Bayle 1880, Journal de Conchyliologie 28, p. 242.

Holopholas (Scobina) costata Linné, Fischer 1887, Manuel de Conchyliologie, Paris, p. 1133.

Pholas (Barnea, section Scobinopholas) costatus Linné, Grant and Gale 1931, Memoirs San Diego Society
Natural History 1, p. 431.

Distinctive characters. Shell oval in outline, rounded at both ends and with strong
radial sculpture extending the entire length. Umbonal reflections well separated from the
surface of the umbos. Protoplax triangular in outline and largely chitinous. Mesoplax
calcareous, transverse and very solid. Apophyses large, broadly spoon-shaped and hollow
at the upper end.

Description. Shell oval in outline, reaching about 183 mm. (about 74 inches) in length
and 75 mm. (about 3 inches) in height, gibbose, rather light but strong, rounded at both
ends, with a long narrow pedal gape, and with a strong radial sculpture covering nearly
the entire shell. Color a chalky-white, with occasional specimens having irregular bands
and markings of pink. Umbos prominent, rather broad, partially covered by the umbo-
nal reflections and located near the anterior fourth of the shell. Umbonal reflections
raised well above the surface of the umbos and having a single support at the posterior
margin with sockets for the reception of the anterior projections of the mesoplax. Sculp-
ture consisting of relatively weak concentric ridges and very strong radial ribs which exist
over the entire length of the shell. Imbrications are formed where the concentric ridges

36 JOHNSONIA, No. 33 Cyrtopleura

and radial ribs cross one another. These imbrications are strong on the anterior and pos-
terior slopes but are slightly reduced over the disc. Numerous fine growth lines are visi-
ble between the concentric ridges. Interior of the shell white and glazed. On occasional
specimens there may be irregular bands and markings of pink. The external sculpture is
clearly visible internally, giving the inside of the valve a ribbed and pitted appearance.
Pallial sinus not apparent, but anterior and posterior adductor muscle scars are fairly well
marked. Occasional specimens have a ledge built out below the posterior adductor mus-
cle scar giving more area for the attachment of the muscle. Apophyses short, wide,
broadly spoon-shaped and hollow at the end. They are marked with concentric growth
lines and occasionally with longitudinal ridges. Protoplax large, thin, triangular in out-
line, composed largely of chitin, though in older specimens it is usually impregnated with
a small amount of calcium. It is pointed and grooved anteriorly, broad and occasionally
slightly lobed posteriorly. It has a central nucleus and faint to rather strong growth lines.
Mesoplax transverse, more or less triangular in outline, solid, calcareous and in one piece.
Periostracum thin, light straw-yellow and deciduous.

Siphons united, smooth, devoid of periostracum, grayish-white to light-ivory in color
with an occasional specimen having a band of buff at the distal end. Incurrent siphon
slightly shorter than the excurrent siphon but with a much larger aperture. Internally
the incurrent siphon has two large dorsal ridges and six to eight small ridges which ex-
tend the entire length of the siphon. These ridges terminate at the opening in minute
cirri. The area between the ridges is marked with mahogany-red vermiculations. Excur-
rent siphon edged with brown and colored for a short distance internally with a hght
straw-yellow. It has one large ventral ridge and numerous smaller ridges which extend
but a short distance anteriorly. Mantle and foot white to light-ivory. Foot elliptical in
outline and truncate.

length width ratio h:]

183.0mm. 72.0mm. 2.54 Mount Hope Bay, Fall River, Massachusetts
165.5 58.5 2.82 Cedar Keys, Florida

147.5 50.0 2.95 Bradenton Beach, Florida

140.0 55.5 ae Clam Bayou, Sanibel Id., Florida

100.5 36.0 OF 82 Galveston, Texas

Types. According to Hanley 1855, Ipsa Linnaei Conchylia, p. 24, Linné did not have
a specimen of C. costata in his collection. His only reference was to Gualtieri 1742, Index
Testarum Conchyliorum, plate 105, fig. G. We select this figure to represent the type,
as it is on the basis of this figure that the species has been well understood. Linné’s orig-
inal locality of Western Europe was corrected to America by Gmelin in the 13th edition
of the Systema Naturae in 1790. We here restrict the type locality to Charleston, South
Carolina, a locality from which we have a good series and one from which specimens may
have reached Europe at that early date.

Remarks. Cyrtopleura costata Linné is one of the largest, most easily recognized, and
best known species in the Pholadidae. Adult specimens can be readily differentiated from
all other species by their large size and by the coarse sculpture which covers the entire
shell. Young specimens might be confused with C. lanceolata d’Orbigny, a much smaller
species. However, this latter species has a much weaker sculpture and the radial ribs are
completely lacking on the posterior slope. From C. cruciger Sowerby it differs by being

Western Atlantic JOHNSONIA, No. 33 37

rounded rather than beaked anteriorly and by having strong radial ribs on the posterior
slope.

Cyrtopleura costata Linné has a wide distribution throughout the Western Atlantic
and can be very abundant in certain restricted areas. These clams live in sandy mud at and
just below low water mark in protected areas, or well below the low tide line on exposed
outer beaches. ‘They can burrow to a depth of two feet or more in the mud and are cap-
able of moving up and down in their burrows at will. When undisturbed, the siphons
are usually seen extending a short distance out of the burrow.

On the west coast of Florida, in an area extending from around Bradenton south to
Englewood Beach, there are a number of colonies of C. costata in which many of the
specimens are variously marked with a bright pink. This pink coloration may be in a
broad band around the umbonal area or in a band near the ventral margin. In some
specimens it is restricted to the posterior slope and in a few the protoplax may be pink.
To date there is no explanation for this rather unusual expression of color in a family
which is otherwise practically devoid of color.

Angel Wings, as this species is popularly called, are one of our most beautiful bivalves
and are much sought after by collectors. They are considered an excellent food and have
appeared on the markets, particularly in Cuba, though not recently.

Capulus shreevei Conrad, described from Long Island, South Carolina beach drift, ap-
pears to be nothing more than the apophysis of this species.

Dall 1889, Proceedings of the Academy of Natural Sciences Philadelphia, p. 274,

Plate 17. Dorsal plates of Cyrtopleura. Figs. 1-3. Cyrtopleura lanceolata d°’Orbigny, off Cabo Polonio, Uru-
guay. Fig. 1. Dorsal view of umbonal region of opposed valves to show the divided mesoplax in place. Fig.
2. Dorsal view of the right half of the mesoplax. Fig. 3. Dorsal view of the thin chitinous protoplax.
Figs. 4-5. Cyrtopleura costata Linné, Sanibel Island, Florida. Fig. 4. Dorsal view of heavy calcareous meso-
plax. Fig. 5. Dorsal view of the flat, largely chitinous protoplax. The stippled portions indicate areas of
calcification. Figs. 6-7. Cyrtopleura cruciger Sowerby, Pacific coast of Panama. Fig. 6. Dorsal view of meso-
plax. Fig. 7. Dorsal view of the very thin and entirely chitinous protoplax.

$8 JOHNSONIA, No. 33 Cyrtopleura

gives a brief account of the gross anatomy of C. costata, and Kellogg (1915) discusses
the ciliary mechanism.

Range. From Fall River, Massachusetts south through the West Indies to Rio de
Janeiro, Brasil.

Plate 18. Cyrtopleura costata Linné, Bradenton, Florida. A specimen showing the band of pink color (natural

size).

Specimens examined. MassacuvuseEtrts: Fall River (J. Miller); New Bedford (MCZ;
ANSP). New Yorks: Long Beach Bay, Orient, Long Island (R. Latham). New JEr-
sEY: Great Egg Harbor (USNM); Cape May (MCZ; ANSP); Sea Isle City (ANSP).
DeLaware: off New England Creek, Delaware Bay in 4 fathoms (USNM); Rehoboth
Beach (ANSP). Marynanp: off Wolf Trap Light, Chesapeake Bay in 94 fathoms
(USNM). Vireinia: Bryants Point, Severn River; Isaac’s Island, Northampton Coun-
ty (both USNM); 10 miles south of Virginia Beach (MCZ); Buckroe Beach, Chesa-
peake Bay (USNM; ANSP); Hog Island, Eastern Shores (USNM). Norru Caro.uina:
Fort Macon, Beaufort ; Long Beach, near South Port (both USN M); Cape Fear(ANSP).
SourH Caroutina: Myrtle Beach (ANSP); South Island, Georgetown County; Cape
Romain; Isle of Palms; Folly Island; Bird Key (all Charleston Museum); Murrell’s
Inlet (A. Merrill); Bull’s Island (J. L. Chamberlin); Sullivan’s Island, Charleston
(Charleston Mus.; MCZ; AMNH; USNM); Beaufort (MWCZ; USNM); Ashe Island,
Edisto River (ANSP). Georera: St. Simons Island (MCZ; ANSP); Sea Island, St.
Simons Island (USNM: ANSP): Savannah (USNM). FrLoripa: Atlantic Beach, near

Western Atlantic JOHNSONIA, No. 33 39

Jacksonville (MCZ); St. Augustine (Univ. of Michigan; USNM: ANSP); Anastasia
Island (ANSP); Daytona Beach (Univ. of Miami); Cape Canaveral; Cocoa Beach (both
MCZ); Cape Sable (MCZ; N.E.Schmidt); Pavilion Key (MCZ); Naples; Caxambas
Pass; Bunch Beach; Kice Island; Marco Island; Lemon Bay; Sarasota: Lossmans Key
(all N. E. Schmidt); Sanibel Island (MCZ; USNM; ANSP); Punta Rassa(AMNH):
Venice (A. Cahn): Bradenton (M. Teare;: USNM): St. Petersburg (Univ. of Michigan:
USNM: MCZ); Terra Ceia; Bonita Beach; Longboat Key; Sarasota (all USNM);
Apalachicola Bay; Fort Myers Beach; Cape Romano (all ANSP); James Island, Frank-
lin County (T. Pulley); Cedar Keys (USNM; ANSP). Atasama: Mobile Bay (MCZ);
Fort Morgan; Little Lagoon, Gulf Shores (both H. I. Johnstone). Mississippi: Long
Beach (ANSP); Deer Island, Biloxi (MCZ). Louisiana: Calasten Pass; Grand Lake
(both USNM); Grand Isle (MCZ). Texas: Matagorda Bay; Keller Bay; Carancahua
Bay (all USNM); 25 miles south of Port Arthur; near Corpus Christi (both MCZ);
Galveston; High Island, Bolivar Peninsula; Boca Chica, near Brownsville (all TT. Pul-
ley); Gulf Beach, Port Isabel (L. A. Weisenhaus). Mexico: Tuxpam; Isla del Carmen,
Campeche (both M. Bourgeois); Tampico, Tamaulipas; Boquilla de Piedras; Tecolutla;
Nautla; Vera Cruz; Alvarado, all of the last four in the State of Vera Cruz (all T. Pul-
ley). Cusa: Habana(MCZ); Mariel, Pinar del Rio (C.G. Aguayo). Hispanioua: Monte
Cristi, Santo Domingo (MCZ). Purerro Rico: Rio Herrera near Loisa Vieja; Ponce
(both MCZ). Lesser AnvILLES: Moruga, Trinidad (H.G. Kugler). British Guiana:
4 miles east of Georgetown (H. G. Kugler). Durcn Guiana: Matappica (C. Bayer).
BrasiL: Ilha de Itaparica, Bahia; Ilha do Cardoso, Cananea, Sio Paulo (both de Oliv-
eira); Nova Almeida (MCZ); Aracaju, Sergipe: Parada Sahy, Sepetiba Bay, Rio de
Janeiro (both H. S. Lopes); Rio de Janeiro (USN M).

Cyrtopleura (Scobinopholas) lanceolata d@ Orbigny
Plates 17 and 19
Pholas lanceolata d’Orbigny 1846, Voyage Amérique Méridionale 5, pt.3, Mollusques, p. 497, pl. 77, figs.
18-19 (L’Ensenada de Ros au Sud du Rio Negro, Patagonie).
Barnea lanceolata d’Orbigny, Tryon 1862, Proc. Academy Natural Sciences Philadelphia 14, p. 208: Car-
celles 1944, Revista del Museo de la Plata (n.s.) Zoologia 3, p. 295, pl. 14, fig. 110.

Distinctive characters. Shell lanceolate in outline and rounded at both ends. Radial
sculpture weak and usually entirely lacking on the posterior slope. Protoplax ‘T-shaped,

Plate 19. Cyrtopleura lanceolata d’Orbigny. Figs. 1-2. Monte Hermoso, Bahia Blanca, Buenos Aires, Argen-
tina (natural size). Fig. 3. Mar del Plata, Buenos Aires, Argentina (hinge area enlarged to show apophysis).

40 JOHNSONIA, No. 33 Cyrtopleura

entirely chitinous and very thin. Mesoplax in two pieces, calcareous and solid. Apophy-
ses short, broad and flattened at the free end.

Description. Shell lanceolate in outline, reaching 69 mm. (about 2# inches) in length
and 21.5 mm. (about £ inch) in height, thin, rounded to slightly angled anteriorly and
narrowly rounded posteriorly. Pedal gape long and narrow, posterior gape slight. Color
a dull chalky-white. Umbos prominent and located near the anterior third of the shell.
Umbonal reflections well separated from the umbos, and with posterior supports which
have sockets for the reception of the mesoplax. Sculpture consisting of well marked con-
centric ridges and rather weak radial ribs which are indicated mainly by the slight imbri-
cations at the point where the radial ribs and concentric ridges cross. Radial ribs are
lacking on the posterior slope in adult specimens but the concentric ridges are well marked.
In young specimens, however, well developed imbrications are evident the entire length
of the shell. Interior of the shell white and glazed. Pallial sinus wide and deep, extend-
ing inward about two thirds the distance to the umbo. Pallial line and muscle scars clearly
indicated in older specimens. Apophyses short, broad, rather thin, fragile and flattened
at the free end. Protoplax exceedingly thin, entirely chitinous and T-shaped. This pro-
toplax is so thin and so closely attached to the muscle that it is easily overlooked. It
curves slightly downward at its posterior extremity, has a posterior nucleus and faint
growth lines which are visible when viewed with transmitted light. Mesoplax transverse,
‘alcareous and in two parts. The anterior extensions of the mesoplax fit into sockets
formed by the folds of the posterior supports of the umbonal reflections. Periostracum
thin, light straw-yellow and deciduous.

Siphons united and covered by a light-brown, minutely papillose, chitinous sheath.
Incurrent siphon with seven to eight large and several small, branched cirri surrounding
the aperture. The excurrent siphon lacks cirri. Pedal gape and foot elliptical in outline.
Foot and mantle white to light-ivory in color.

length width ratio h:]
69.0 mm. 21.5 mm. 8.2 Monte Hermoso, Bahia Blanea, Buenos Aires, Argentina
56.5 18.0 3.1 Playa de Punta Medenas, Buenos Aires, Argentina

Types. The holotype of Pholas lanceolata d’Orbigny isin the British Museum, accord-
ing to Gray 1854, List of the Shells of South America in the Collection of the British
Museum Collected and Described by M. Alcide d’Orbigny, p. 55. The type locality is
Ensenada de Ros south of Rio Negro, Patagonia, Argentina.

Remarks. Cyrtopleura lanceolata d’Orbigny differs from C. cruciger Sowerby, with
which it might be confused, by having the anterior end of the valves rounded rather than
moderately beaked, by being lanceolate rather than oval in outline, by having a much
weaker sculpture and by having the mesoplax in two pieces. From young specimens of
C. costata Linné it may be distinguished by its weak radial sculpture which is lacking on
the posterior slope, by its flat rather than spoon-shaped apophyses, by its proportion-
ately longer anterior slope, and also by having the mesoplax in two pieces.

We know nothing of the biology of this species. It has seldom been recorded in the
literature and there are but few specimens in the museums of this country. However,
through the kindness of A. Carcelles, formerly of the Museo Nacional de Argentina, we
have had a fine series to study.

Range. From Santos, Brasil south to the Gulf of San Matias, Argentina.

Western Atlantic JOHNSONIA, No. 33 11

Specimens examined. Brasit: Santos (USNM);: Frances Island, north of Santa Cath-
arina Island, Estado Santa Catharina (H. Lopes). UruGuay: off Cabo Polonio (S. Lat.
34°42’; W. Long. 54°10’) (A. Carcelles). ArGENTINA: Mar del Plata, Buenos Aires;
Monte Hermoso, Bahia Blanca, Buenos Aires; Playa de Punta Medanos, Rio Negro
(all A. Carcelles): Buenos Aires (USNM); Carmen de Patagones, Buenos Aires (Univ.
of Michigan: USNM): Rio Negro (Redpath Museum, Montreal; USNM).

Subgenus Cyrtopleura Tryon
Cyrtopleura Tryon 1862, Proceedings Academy Natural Sciences Philadelphia 14, p. 201.

Shell beaked anteriorly, rounded posteriorly, and having a broad oval pedal gape. Ac-
cessory plates consisting of a more or less T-shaped, chitinous protoplax and a transverse,
calcareous mesoplax. Umbonal reflections well separated from the umbos and supported
at the posterior margins where they curve downward and form sockets for the reception
of the anterior extensions of the mesoplax. Apophyses broad and flattened.

Subgenotype, Pholas crucifera Sowerby (= Pholas cruciger Sowerby), subsequent des-
ignation, Stoliczka 1870.

Cyrtopleura (Cyrtopleura) cruciger Sowerby
Plates 17, 20 and 21

Pholas cruciger Sowerby 1834, Proceedings Zoological Society London, p. 69 (Puna Island in Gulf of Guay-
aquil and Bay of Caraccas [Caraques], West Colombia [Ecuador] and in the Gulf of Nocoiyo [Nicoya], Costa
Rica).

Pholas crucifera Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, p. 489 [error for cruciger Sowerby ].

Pholas crucigera Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, pl. 104, figs. 24-26 error for cruciger,
on plate caption only].

Pholas (Cyrtopleura) crucifera Sowerby, Tryon 1862, Proceedings Academy Natural Sciences Philadelphia,
14, p. 201.

Cyrtopleura exilis Tryon 1870, American Journal of Conchology 5, p. 170, pl. 14, fig. 2 (Ins. St. Croix
[Virgin Islands], West Indies).

Plate 20. Cyrtopleura exilis Tryon (=cruciger Sowerby) St. Croix, Virgin Islands. Lectotype, ANSP 51040
(about 3x).

42 JOHNSONIA, No. 33 Cyrtopleura

Distinctive characters. Shell beaked anteriorly, narrowly rounded posteriorly and with
concentric sculpture extending over the entire surface of the shell. Radial sculpture strong
on the anterior slope, becoming weak on the disc and lacking on the posterior slope. Ac-
cessory plates consisting of a very thin, chitinous, more or less ‘T-shaped protoplax and a
transverse mesoplax.

Description. Shell oval in outline, reaching about 44 mm. (1# inches) in length and 18
mm. (about 2 inch) in height, thin but strong, rounded posteriorly and with small rounded
beaks anteriorly. Pedal gape large, generally oval in outline and tapering at both ends.
Color a dull chalky-white to light-brown. Umbos prominent and located near the an-
terior third of the shell. Umbonal reflections well separated from the umbos and with a
single support at the posterior margins where sockets are formed for the reception of the
anterior projections of the mesoplax. Posterior to the umbo, the dorsal margin of the
valve is reflected upward and outward forming a sharp angle with the margin of the um-
bonal reflection. Sculpture consisting of rather strong concentric ridges and radial ribs.
Concentric ridges are well marked throughout the length of the shell, while the radial
ribs are strong on the anterior slope, weak on the disc, and lacking on the posterior slope.
Imbrications are formed where the concentric ridges and radial ribs cross one another.
These imbrications are prominent on young and perfect specimens, but on worn adult
specimens they may be indicated only as slightly roughened wavy lines. Sculpture below
the umbonal reflection consisting of crowded concentric ridges. Interior of shell white
and glazed and with the radial ribs clearly showing on the anterior portion. Pallial sinus
broad and deep, extending inward about the distance to the umbo. Apophyses short,
broadened and flattened, curved sharply inward and solid at the upper end. Protoplax
very thin, entirely chitinous and generally more or less ‘T-shaped. Mesoplax transverse,
strong, calcareous and in one piece. Periostracum exceedingly thin, light straw-yellow
and deciduous.

The siphons may be extended 8 to 4 times the length of the shell; they are light-tan
in color and minutely papillose. Incurrent siphon with 6 large and numerous small
branched cirri. Foot elliptical in outline, truncate and white to light-ivory in color.
Mantle white to light-ivory. The above description of the soft parts is based upon pre-
served material.

length height ratio h:]
36.0 mm. 17.5 mm. 2.0 Panama
44.0 18.0 2.4 -
38.7 16.5 2.3 -:

Types. The types of Pholas cruciger Sowerby are in the British Museum. Sowerby
listed three localities in his original description: Puna Island in the Gulf of Guayaquil
and the Bay of Caraccas [Caraques], Colombia [Ecuador] and in the Gulf of Nocoiyo
[Nicoya], Costa Rica. We here restrict the type locality to the Gulf of Nicoya, Costa
Rica, a locality from which we have seen specimens. The lectotype of Cyrtopleura evilis
Tryon is inthe Academy of Natural Sciences of Philadelphia, no. 51040. The type local-
ity of St. Croix, Virgin Islands, we believe, is in error.

Remarks. Cyrtopleura cruciger Sowerby is a very distinctive species not closely related
to any other in this group. It differs from young specimens of C. costata Linné by lack-
ing the radial ribs on the posterior slope, by having the anterior margin of the shell

Western Atlantic JOHNSONIA, No. 33 $3

beaked and by having the apophyses solid at the upper end. ‘There are also considerable
differences in the dorsal plates which can best be seen by studying the figures. From C.
lanceolata d’Orbigny it differs by being oval rather than lanceolate in outline, by having
a much stronger sculpture, by being beaked anteriorly, and by having the mesoplax in
one piece.

Cyrtopleura cruciger Sowerby is apparently a rare species throughout its range and is
restricted in its ecological distribution to areas of soft stone. We have seen several speci-
mens of a soft stone taken at Panama with the shells of C. eruciger still in place. They
generally bore to a depth of about twice the length of the shell and make a rather snug
burrow.

Philippi (1851, Abbildungen und Beschreibungen Conchylien 3, p. 129) was in error
when he gave one of the localities for cruciger as ‘‘sinus Caraccas [sic] in Mari Antilla-
rum.’ Sowerby in his original description stated that this species, which was collected
by Hugh Cuming, came from the “‘island of Puna in the Gulf of Guayaquil; in soft stone
at low water in the Bay of Caraccas; both in West Colombia.’’' This Eastern Pacific spe-
cies is not known from the Western Atlantic. Cyrtopleura evilis Tryon is definitely this
species and the locality St. Croix is certainly in error.

Range. From Guaymas, Sonora, Mexico south to Puna Island in the Gulf of Guay-
aquil, Ecuador.

Specimens examined. Mexico: Guaymas, Sonora (USNM). Cosra Rica: Isla San
Lucas, Gulf of Nicoya (USNM). Panama: (Redpath Museum; C. M. Dumbauld:
MCZ); Panama City (J. Zetek); Old French Canal, Canal Zone (ANSP).

Plate 21. Cyrtopleura cruciger Sowerby. Figs. 1-4. Panama (all natural size). Fig. 1. External view showing
the umbonal reflection with its posterior support and the notch formed by the reflection of the dorsal mar-
gin of the valve posterior to the umbos. Fig. 2. Interior view showing the broad, flat apophysis, the muscle
sears and pallial sinus. Fig. 3. Dorsal view of opposed valves showing the narrow umbonal reflection and
the supports. Fig. 4. Ventral view of opposed valves showing the beaks and the pedal gape.

‘The Bay of Caracas in which Hugh Cuming collected was in Colombia, now Ecuador (South Latitude
° 5 ae) . 2 , ‘ ~ : .
0°30’). This place is now known as Bahia de Caradques. Clench 1945, Occasional Papers on Mollusks 1, p. 26.

44 JOHNSONIA, No. 33 Phelan

Genus Pholas Linné

Pholas Linné 1758, Systema Naturae, ed. 10, 1, p. 669; Lamarck 1799, Prodrome d’une Nouvelle Classi-
fication des Coquilles [in] Mémoires de la Société d’ Histoire Naturelle Paris, p. 90; non Lamarck 1801.

Genotype, Pholas dactylus Linné, subsequent designation, Children 1822.

Shells more or less elliptical in outline, with double, septate umbonal reflections and
three accessory dorsal plates. Protoplax oval to quadrangular in outline, thin, calcareous,
in one part or divided longitudinally into two parts. Mesoplax transverse and more or
less triangular in outline, calcareous and solid in structure. Metaplax long and narrow.
Shell beaked or rounded anteriorly. Sculpture extending over the entire shell or lacking
on the posterior slope.

The genus may be divided into three subgenera on the basis of the above characters
and may be keyed out as follows:

1. Protoplax divided into two parts 2

Protoplax in one piece Monothyra (Indo-Pacific)
2, Shell beaked anteriorly, nuclei of the divided protoplax

located near the posterior outer margin Pholas s.s.

Shell rounded anteriorly, nuclei of the divided protoplax

located near the anterior inner margin Thovana

Subgenus Pholas Linné

Pholas Linné 1758, Systema Naturae, edition 10, 1, p. 669; Lamarck 1799, Prodrome d’une Nouvelle
Classification des Coquilles [in] Mémoires de la Société d’ Histoire Naturelle Paris, p. 90; non Lamarck 1801.

Hypogaea Poli 1791, Testacea Utriusque Siciliae Eorumque Historia et Anatome 1, Introduction, p. 29
[name for soft parts only ].

Hypogaeoderma Poli 1795, Testacea Utriusque Siciliae Eorumque Historia et Anatome 2, pp. 251, 257 (type,
here selected, Pholas dactylus Linné).

Dactylina Gray 1847, Proceedings Zoological Society London, p. 187 (genotype, Pholas dactylus Linné,
monotypic); non Dactylina Zborzewski 1843.

Xylotrya ‘Leach’ Menke 1830, Synopsis Methodica Molluscorum, p. 121 (genotype, Pholas dactylus Linné,
subsequent designation, Clench and Turner 1946).

Pragmopholas Fischer 1887, Manuel de Conchyliologie, p. 1133 (genotype, here selected, Pholas dactylus
Linné).

Phragmopholas ‘Fischer’ Dall 1898, Transactions Wagner Free Institute of Science, Philadelphia, 3, p.
814 [error for Pragmopholas Fischer}.

Subgenotype, Pholas dactylus Linné, subsequent designation, Children 1822.

The subgenus Pholas is characterized by having the shells beaked anteriorly, by having
the protoplax divided longitudinally into two parts, and by having the nuclei of the di-
vided protoplax near the posterior outer margin.

This subgenus is restricted in its distribution to the Eastern Atlantic. The following
description of Pholas dactylus Linné, the genotype of Pholas, is included for the clarifi-
cation of the genus.

Western Atlantic JOHNSONIA, No. 33 45

Pholas (Pholas) dactylus Linné

Plates 22 and 23

Pholas dactylus Linné 1758, Systema Naturae ed. 10, 1, p. 669 (Europe); Bucquoy, Dautzenberg and Doll-
fus 1896, Mollusques Marins du Roussillon 2, p. 609, pl. 87.

Pholas muricatus DaCosta 1778, Historia Naturalis Testaceorum Britanniae, p. 244, pl. 16, fig. 2 (shores of
Great Britain and Ireland).

Pholas dactilus “Linné’ Born 1778, Index Rerum Naturalium Musei Caesarei Vindobonensis, p. 7 [error for
dactylus Linné].

Hypogaea verrucosa Poli 1795, Testacea Utriusque Siciliae Eorumque Historia et Anatome 2, p. 251 [name
given to the soft parts only].

Hypogaeoderma dactylus Linné, Poli 1795, Testacea Utriusque Siciliae Eorumque Historia et Anatome 2,
De 257.

Pholas hians Solander 1786, Catalogue of the Portland Museum, p. 102, lot 2240; p. 174, lot 3736 [nomen
nudum |.

Pholas hians ‘Solander’ Pulteney 1799, Catalogue of the Birds, Shells, ete. of Dorsetshire, p. 26 [in]
Hutchins’ History of the County, Dorset, London. (Dorset coast at Waymouth [sic], Swanage and the north
shore at Poole); non Pholas hians Gmelin 1790.

Pholas callosa Lamarck 1818, Animaux Sans Vertébres 5, p. 445 (Bayonne [France] ); Lamy 1922, Bulle-
tin Museum National d’Histoire Naturelle, Paris, 28, p. 245.

Dactylina dactylus Linné, Gray 1847, Proceedings Zoological Society London, p. 187.

Pholas dactylus var. gracilis Jeffreys 1865, British Conchology 3, p. 105 (Exmouth, England).

Pholas dactylus var. decurtata Jeffreys 1865, British Conchology 3, p. 105 (Sussex, England).

Pholas dactylina Locard 1886, Prodrome de Malacologie Francaise, Catalogue Général des Mollusques Vi-
vantes de France. Mollusques Marins, p. 365 [in foot note].

Pragmopholas (Dactylina) dactylus Linné, Fischer 1887, Manuel de Conchyliologie, p. 1133, fig. 863.

Distinctive characters. Shell white, beaked anteriorly, rounded posteriorly, with
double, septate umbonal reflections and having a divided protoplax, a triangular meso-

Plate 22. Pholas dactylus Linné. Figs. 1-2. Island of Malta (slightly reduced). Fig. 1. Internal view showing
the broad flat apophysis, muscle scars and deep pallial sinus. Fig. 2. External view showing the septate
umbonal reflection, the beak, and the pronounced concentrie sculpture.

46 JOHNSONIA, No. 33 Pholas

plax, and a long narrow metaplax. Nuclei of the divided protoplax located near the outer
posterior margin.

Description. Shell subelliptical in outline, reaching about 130 mm. (about 5 inches) in
length and 41 mm. (about 14 inches) in height. Shell thin but strong, beaked anteriorly,
rounded posteriorly, with septate umbonal reflections. The sculpture is often weak
or absent on the posterior slope. Color a dull chalky-white to grayish-white. Umbos
prominent, located near the anterior fourth of the shell and covered by double septate
umbonal reflections. There are 10 to 14 septa in the average specimen about 3 inches
long. Umbonal reflections free for a short distance over the beaks but closely appressed
over the umbos and posterior to them. Sculpture consisting of laminated concentric
ridges and radial ribs. Laminated ridges well marked for the entire length of the shell in
young specimens, but in older specimens and especially those boring into hard rock they
may be entirely lacking on the posterior slope. Radial ribs prominent on the anterior
slope but becoming weak over the dise and entirely lacking on the posterior slope. Strong
imbricated scales are produced where the concentric ridges and radial ribs cross one an-
other. However, these may be worn down to undulating ridges in specimens boring into
a hard substratum. In young specimens the concentric ridges are rather widely spaced
and prominent the entire length of the shell. Very young specimens may be imbricate
even on the posterior slope. Interior of the shell white and glazed witha slight indication
of the radial ribs and concentric ridges showing through. Umbonal reflections usually
worn away at the point of contact and rotation of the two valves. Muscle scars and pal-
lial sinus well marked. Pallial sinus broad and deep, extending inwards two thirds the
distance to the umbos. Apophyses rather short, solid and strong, flattened and often
ridged on the free ends. Accessory plates three, a divided protoplax, a transverse meso-
plax, and a long narrow metaplax. Protoplax calcareous, fragile, divided through the
middle longitudinally, truncate posteriorly, acuminate anteriorly, with well marked con-
centric growth lines and with the nuclei near the posterior outer margins. Mesoplax, as
shown in plate 23, calcareous and solid. Metaplax long and narrow, rather thin and
faintly marked with growth lines.

5

Plate 23. The dorsal plates of Pholas dactylus Linné. Fig. 1. Nantes, France. Dorsal view of an entire speci-
men to show the normal position and arrangement of the dorsal plates. Figs. 2-5. St. Juliac, Dinard, France.
Individual plates of a young specimen. Fig. 2. Dorsal view of the divided protoplax showing the faint growth
lines and the location of the nuclei near the posterior outer margins. Fig. 3. Dorsal view of the mesoplax
tipped forward to show the basal flange. Fig. 4. Mesoplax in normal position. In young specimens this has
a deeply curved posterior margin, while in older specimens it is nearly straight as shown in Fig. 1. Fig. 5.
Dorsal view of the metaplax showing the fine growth lines.

Western Atlantic JOHNSONIA, No. 33 47

— 2

Siphons extending at least one to two times the length of the shell, white to light-ivory
in color, papillose and devoid of periostracum except for a small band near the posterior
margin of the shell. Incurrent siphon nearly twice the diameter of the excurrent siphon
and with 10 to 14 small-branched cirri surrounding the aperture. Excurrent siphon lack-
ing cirri. We have seen only preserved specimens of this species but the gross morphology
has been figured by Poli, Deshayes, and DuBois.

length height ratio h:]

149.5mm. 41.5 mm 3.5 Malta (fish market)

132.0 43.0 3.1 Malta

113.0 33.0 3.4 Smyrna, Turkey

105.0 32.0 3.8 + >
94.0 32.0 2 Cornwall, England
81.0 28.5 2.8 Pouliguen, near Nantes, France
28.0 14.0 2.0 La Rochelle, France

Types. According to Hanley (1855, [psa Linnaei Conchylia, p. 24) the types of Pholas
dactylus are in the Linnean Collection. The type locality given by Linné was Europe.
The location of the type of Pholas muricatus DaCosta is unknown. The types of P.
hians Pulteney, P. dactylus var. gracilis Jeffreys and P. dactylus var. decurtata Jeffreys
are probably in the British Museum.

Remarks. Pholas dactylus Linné is a very variable species both in its appearance and
in its habitat. The substrata in which it has been found boring vary from such easily
worked material as sand, peat, marl, and waterlogged wood to slate, sandstone and schis-
tose rocks. Jeffreys states that his variety gracilis was boring in pure sand and as a con-
sequence it was a thin and delicate shell. His decurtata, which he described as stunted
and solid with close and usually effaced sculpture, was found in hard rocks. It is this
wide range of habitat that has been responsible for these ecological forms and has resulted
in the extensive synonymy given above.

Pholas dactylus Linné is perhaps the most famous of all pholads. It was well known
to the early naturalists of Europe who were fascinated by its luminescent properties and
its ability to bore into hard rock. Much has been written on this and a summary of some
of the work is given in the introduction to this paper. In the early Mediterranean coun-
tries Pholas dactylus Linné was considered a great delicacy and it is still eaten by some
today.

Range. From the Firth of Forth, Scotland (Hanley 1853, p. 11) south along the East-
ern Atlantic coast to Oran, Algeria; the Mediterranean, Adriatic, and Aegean Seas.

Specimens examined. Brirish Istes: Tenby, Wales; Cornwall, South Devon; Kent
County (all MCZ); Broadstairs (Charleston Museum). FRANCE: Caleu, near La Roch-
elle; coast of Pouliguen, near Nantes; Nantes; Rouen: St. Juliac, Dinard (all MCZ);
La Rochelle; Touraine (both USNM). Iraty: Porto Maurizio; Gulf of Naples (both
MCZ). Mauresr Istanps: Senglea Point, Malta (MCZ). Turkey: Smyrna (MCZ).
ALGERIA: Oran (MCZ).

48 JOHNSONIA, No. 33 Pholas

Subgenus Thovana Gray

Thovana ‘Leach’ Gray 1847, Proceedings Zoological Society London, p. 187 (subgenotype, Pholas oblongata
Say, monotypic).

Gitocentrum Tryon 1862, Proceedings Academy Natural Sciences Philadelphia 14, pp. 203-204 (subgeno-
type, here selected, Pholas campechiensis Gmelin).

Shell rounded anteriorly, with septate umbonal reflections and three accessory plates.
Protoplax divided into two parts longitudinally with the nuclei anterior and more or less
centrally located. Mesoplax small and transverse. Metaplax thin, long and narrow.

The subgenus Thovana is restricted in its distribution to the Western Atlantic and
the Eastern Pacific.

Subgenotype, Pholas oblongata Say (=P. campechiensis Gmelin), monotypic.

Pholas (Thovana) campechiensis Gmelin
Plates 24 and 25

Pholas campechiensis Gmelin 1790, Systema Naturae, edition 13, 1, p. 3216 (Sinu Campechiensi).

Pholas oblongata Say 1822, Journal Academy Natural Sciences Philadelphia 2, p. 320 (Georgia, Carolina
and East Florida); non oblongata Tuomey and Holmes 1858.

Thovana oblongata Say, Gray 1847, Proceedings Zoological Society London, p. 187.

Dactylina (Gitocentrum) campechensis ‘Gmelin’ Tryon 1862, Proceedings Academy Natural Sciences Phila-
delphia 14, p. 204 [error for campechiensis Gmelin].

Pholas candeana d’Orbigny 1842 [in] Sagra, Histoire de L’Ile de Cuba, Mollusques 2, p. 215, pl. 25, figs.
18-19; (Martinique; Habana, Cuba; Florida).

Pragmopholas (Gitocentrum) campechiensis Gmelin, Fischer 1887, Manuel de Conchyliologie, p. 11338.

Distinctive characters. Shell white, rounded at both ends, with septate umbonal reflec-
tions, with the concentric sculpture extending the entire length of the shell and with
three accessory plates consisting of a divided protoplax, a mesoplax and a metaplax.

Description. Shells subelliptical in outline, reaching about 110 mm. (about 42 inches)
in length and 85 mm. (12 inches) in height, thin, fragile, rounded, gaping slightly at
both ends, and with the sculpture extending over the entire shell. Color a chalky-white
to gray-white. Umbos prominent, located near the anterior fourth of the shell and cov-
ered by double, septate umbonal reflections. There are 10 to 14 septa in the average
specimen of about 3 inches in length. Umbonal reflections free anterior to the umbos,
but closely appressed over the umbos and posterior to them. Sculpture consisting of
laminated, concentric ridges and radial ribs. Concentric ridges strong on the anterior
slope, becoming somewhat weaker over the disc, and in adult specimens, usually weak
and occasionally absent on the posterior slope. Radial ribs prominent on the anterior
slope, becoming weak on the disc and usually disappearing on the posterior slope. Im-
bricated scales are formed where the concentric ridges and radial ribs cross one another.
In young specimens the concentric ridges are widely spaced, very prominent and imbri-
cate, even on the posterior slope. Sculpture below the umbonal reflections consisting of
low, smooth, concentric ridges. Interior of shell white and glazed. Radial ribs and con-
centric ridges usually visible internally, especially on the anterior slope. Umbonal reflec-
tions usually worn away at the point of contact of the two valves. Muscle scars and pal-
lial line well marked. Pallial sinus broad, deep and extending inward nearly two thirds

Western Atlantic JOHNSONIA, No. 33 19

the distance to the umbo. Apophyses fragile, short and broad, and projecting from be-
neath the umbo at a sharp angle posteriorly. Accessory plates three, a double, nearly
rectangular protoplax, a transverse mesoplax, and an elongate narrow metaplax. Proto-
plax divided through the middle longitudinally, truncate posteriorly, and acuminate an-
teriorly. The nuclei of the divided protoplax are anterior and nearly centrally located.
Mesoplax small, and a broad, flattened triangle in outline when seen in position. Meta-
plax thin, long and narrow.
We have not seen live or even preserved specimens of this species.

length height ratio h:]
108.5 mm. 32.5 mm. 3.3 Sea Island, Georgia
95.0 31.0 3.1 Harbor Island Marsh, New Hanover County, North Carolina
70.0 29-0 3.2 10 miles south of Sabine, Texas
69.5 23.5 2.9 La Brea, Trinidad
63.5 19.5 8.2 Sénégal, Africa

Types. The location of the type of Pholas campechiensis Gmelin is unknown. Gmelin
made only one reference and that was to Lister 1770, Historiae Sive Synopsis Methodi-
cae Conchyliorum et Tabularum, Editio Altera, pl. 432, fig. 275. We here select this
figure to represent the type. The type locality is the Gulf of Campeche, Mexico. The
type of Pholas oblongata Say is apparently lost as it is not in the Academy of Natural
Sciences, Philadelphia where most of Say’s types are located. The type of Pholas can-

deana a’ Orbigny from Cuba is in the British Museum according to Gray 1854.

Remarks. Pholas campechiensis Gmelin is most closely related to P. chiloensis Molina
of the Eastern Pacific. It differs from this species by having a smaller, more delicate shell
and by having the sculpture extending over the posterior slope. In P. chiloensis the sculp-
ture is usually reduced to growth lines only on the posterior slope. See also remarks
under P. chiloensis Molina.

Pholas campechiensis Gmelin though apparently not a rare species is very seldom col-
lected alive. All specimens studied were dead and most of them were beach worn. Only

Plate 24. Pholas campechiensis Gmelin. Figs. 1-2. La Brea, Trinidad (about natural size). Fig. 1. Showing
muscle sears, pallial sinus and spoon-shaped apophysis. Fig. 2. Showing the septate umbonal reflection and
sculpture extending over the posterior slope. Fig. 3. Ten miles south of Sabine, Texas, a specimen with
the sculpture becoming very weak on the posterior slope (about natural size). Fig. 4. Sénégal, Africa
(about 14x).

50 JOHNSONIA, No. 33 Pholas

two specimens, both of them young, had the accessory plates. In certain areas these
shells are common in the beach rubble as indicated by the large number of fragments
found in a pint of drift material collected at High Island, Bolivar, Texas by T. E. Pulley.
Fragments of the umbonal areas of at least one hundred specimens were in this small
amount of drift. This portion of the shell is very strong; it is the last to be destroyed by
wave action and consequently accumulates in the beach drift. It seems probable that P.
campechiensis Gmelin, like Panope bitruncata Conrad, lives rather deep in the mud well
below the low tide line, and, being deeply buried, the shells are not brought up by
dredges. When the animals die the shells remain 7m situ, with the result that only a rel-
atively few specimens are washed out and eventually reach shore.

A specimen received from A. S. Merrill was taken from a piece of waterlogged wood
which had washed ashore at Folly Beach, South Carolina. In the same log there were
several specimens of Barnea truncata Say, Petricola pholadiformis Lamarck, and Marte-
sia cuneiformis Say. We have a second piece of waterlogged wood that was taken at
Daytona Beach, Florida and which also contained several specimens of this species. In
both cases the specimens were young but quite normal and did not appear to be stunted
or malformed.

From the material we have had for study the shells of P. campechiensis Gmelin appear
to be far more uniform in shape and sculpture than are most species in this family. This
would, perhaps, indicate a limited type of substratum in which this species can survive,
probably stiff mud and sand.

Range. From North Carolina south through the West Indies to Sepetiba Bay, Rio
de Janeiro, Brasil. EasrERN ATLANTIC: from Dakar, Sénégal south to Liberia.

Plate 25. Dorsal plates of Pholas subgenus Thovana. Figs. 1-5. Pholas (Thovana) campechiensis Gmelin, Folly
Beach, South Carolina. The plates of this very young specimen (31 mm.in length) are probably more
highly sculptured than would be the case with older specimens. Fig. 1. Dorsal view of divided protoplax
showing the nuclei located near the anterior central portion. Fig. 2. Side view of the right half of the pro-
toplax from the outer edge showing the curvature. Fig. 3. Dorsal view of the mesoplax tipped forward to
show the basal flange. Fig. 4. Dorsal view of the mesoplax in normal position (anterior to right). Fig. 5.
Dorsal view of metaplax which is thin and almost entirely chitinous in young specimens. Figs. 6-10. Pholas
(Thovana) chiloensis Molina, Chile. Fig. 6. Dorsal view of divided protoplax showing the nuclei located near
the anterior central portion. Fig. 7. Side view of left half of the protoplax from the outer edge to show the
curvature. Fig. 8. Dorsal view of the mesoplax tipped forward to show the basal flange. Fig. 9. Mesoplax
in normal position (anterior to the right). Fig. 10. Dorsal view of the metaplax which is thin, fragile, and
only partially impregnated with calcium.

Western Atlantic JOHNSONIA, No. 33 51

Specimens examined. Norra Carouina: Harbor Island Marsh, New Hanover Co.
(MCZ). Ocracoke Island (ANSP). Sourn Carorina: Myrtle Beach; Pawleys Island;
Sullivans Island, Charleston (all Charleston Museum); Magnolia Beach (MCZ); Charles-
ton (USNM);: Folly Island (ANSP; A. Merrill). Grtorcra: Sea Island, Glynn Co.
(MCZ; ANSP); St. Simon's Island (USNM). FLoripa: Jacksonville Beach (MCZ):
St. Augustine (Univ. of Michigan; ANSP; USNM); Daytona Beach (G. Quelch);
Cape Canaveral (MCZ): Cape Sable (H. Moore): Sanibel Island (MCZ). Louisiana:
Cameron (USNM). Texas: 10 miles southwest of Sabine (MCZ); Galveston (‘T. E. Pul-
ley; ANSP); Port Aransas (MCZ); Mustang Island (J. Hedgepeth); Gulf Beach, Port
Isabel (B. Weisenhaus): High Island (T. Pulley): Brownsville(USNM). Grearer AN-
TILLES: Jamaica( ANSP). Lesser ANTILLES: La Brea, Trinidad (MCZ): Moruga and
Guayaguagare Beach, Trinidad (both H. J. Kugler). Mexico: Tampico, ‘Tamaulipas:
Boquilla de Piedras; Tecolutla; Nautla and Vera Cruz, all in the State of Vera Cruz
(all T’. Pulley); Tuxpan and Barra de Alvarado, Vera Cruz (both M. E. Bourgeois).
NicaraGua: Wounta Haulover (MCZ). Panama: Colén (MCZ). CoLompia: Carta-
gena (MCZ). Brasix: [Ilha de Itaparica, Estado de Bahia (J. de Oliveira); Barra Seca,
Espirito Santo (Thayer Expedition, MCZ): Aracaju, Estado de Sergipe: Parada Sahy,
Sepetiba Bay, Estado Rio de Janeiro (both H. S. Lopes).

EasteERN ATLANTIC: SENEGAL: Dakar (Inst. Frangais d’ Afrique Noire). Gop Coast:
Accra (MCZ). Liserta: Marshall (MCZ).

Pholas (Thovana) chiloensis Molina
Plates 25, 26 and 27

Pholas chiloensis Molina 1782, Saggio Sulla Storia Naturale de Chili, p. 204 (Archipelago de Chiloe); King
1832, Zoological Journal 5, p. 334; Hupé [in] Gay 1854, Historia de Chile, Zoologia 8, p. 381, Atlas 2,
Mollusks, pl. 8, fig. 3.

Pholas chiloensis var. parva Sowerby 1834, Proceedings Zoological Society London, p. 69 (soft stone at 17
fathoms, Island of Plata, West Colombia [Ecuador], H. Cuming collector).

Pholas laqueata Sowerby 1849 [1850] Proceedings Zoological Society London, p. 161 [nude name]; Sowerby
1849, Thesaurus Conchyliorum 2, pt. 10, p. 486, pl. 103, figs. 19-20 (Isle of Plata, Colombia [Ecuador], H.
Cuming collector).

Pholas (Dactylina) retifer Mérch 1860, Malakozoologische Blatter 7, p. 177 (Realejo, Nicaragua).

Pholas dilecta Pilsbry and Lowe 1932, Proceedings Academy Natural Sciences Philadelphia 84, p. 88, pl.
11, figs. 8-9 (Corinto, Nicaragua).

Distinctive characters. Shell white, rounded at both ends, with septate umbonal reflec-
tions, with the sculpture greatly reduced or lacking on the posterior slope and with three
accessory plates consisting of a divided protoplax, a mesoplax, and a metaplax.

Description. Shells rather large, reaching 123 mm. (about 4¢ inches) in length and 44
mm. (about 14 inches) in height, elliptical to oval in outline, rounded and gaping slightly
at both ends, with double septate umbonal reflections, and generally lacking sculpture
on the posterior slope. Color a dull chalky-white. Umbos prominent and located near
the anterior fourth of the shell. Umbonal reflections free anterior to the umbos, but
closely appressed, double and septate over the umbos. There are 14 to 16 septa in the
average three to four inch specimen. Sculpture consisting of concentric ridges and radial
ribs with imbrications produced where the two cross one another. The concentric ridges
are very prominent and laminated on the anterior slope, much lower and weaker on the

JOHNSONIA, No. 33 Pholas

Or
G
to

disc, and generally evident only as growth lines on the posterior slope. The radial ribs are
strong on the anterior slope, becoming much weaker on the disc and are completely lack-
ing on the posterior slope. In most specimens there is a rather definite line of demarca-
tion between the sculptured disc and the nearly smooth posterior slope. Interior of shell
white and glazed, the external sculpture showing internally as aseries of grooves. Muscle
scars and pallial line well indicated. The pallial sinus is broad and deep, extending inward
three fourths of the distance to the umbo. Apophyses rather heavy, slightly enlarged at
the free end, and extending outward from under the umbo at a sharp angle posteriorly.
Umbonal reflections usually worn down at the point of contact of the two valves. Proto-
plax divided longitudinally into two parts. Each half is truncate posteriorly, acuminate
anteriorly, has well marked growth lines and anterior nuclei which are located near the
inner margin. Mesoplax transverse and nearly diamond-shape in outline. Metaplax long
and narrow, exceedingly fragile and only slightly to moderately impregnated with cal-
cium. Periostracum thin, light straw-yellow, and deciduous.

Siphons a pale brown in color (an old preserved specimen), separated slightly at their
posterior extremity and with small uniform papillae covering the entire surface. Foot
elliptical in outline, long and thin. Foot and mantle light-ivory in color.

length height ratio h:]
122.5mm. 39.0mm. 8.1 Kino Bay, Sonora, Mexico
112.5 37.5 3.0 » ar a3 *
114.5 45.5 a5 Paracas, Peru
93.5 29.0 3.2 Panama City, Panama
90.0 30.0 3.0 Chiloe Island, Chile
15.0 22.0 3.4 Holotype, P. di/ecta Pilsbry and Lowe
(Ee DRE: 3.2 Cotype, P. laqueata Sowerby

Plate 26. Pholas chiloensis Molina. Figs. 1-2. Chiloe Island, Chile. Fig. 3. Holotype of Pholas dilecta Pilsbry
and Lowe (=chiloensis Molina) from Corinto, Nicaragua. This specimen was not photographed in the con-
ventional position but was tipped forward in order to show the remains of the septate umbonal reflection,
most of which had been broken off. In this position the shell appears to be narrower than it is (all natural
size).

Western Atlantic JOHNSONIA, No. 33 53

Types. The location of the type specimens of Pholas chiloensis Molina is unknown.
We here select the figure given by Hupé in Gay 1854, Historia de Chile, Zoologia 8,
p. 881, Atlas 2, pl. 8, fig. 3 to represent the type. The type locality is Chiloe Island,
Chile. We figure a topotype. The type specimen of P. chiloensis parva Sowerby from
the Island of Plata, Ecuador and P. laqueata Sowerby from the same locality are appar-
ently lost. They are not in the Cuming collection in the British Museum according to a
letter received from G. L. Wilkins. We have seen a probable cotype of the latter species,
originally from the Cuming collection and now in the Zoologische Museum, Amsterdam.
The whereabouts of the types of Pholas retifer Mérch is unknown. The holotype of
Pholas dilecta Pilsbry and Lowe from Corinto, Nicaragua is in the Academy of Natural
Sciences Philadelphia, no. 155299.

Remarks. Pholas chiloensis Molina is very closely related to P. campechiensis Gmelin
of the Western Atlantic. Young specimens of the two are difficult or even impossible to
distinguish. In older specimens, P. chiloensis Molina can be distinguished by its larger
size, heavier shell, its generally higher umbonal reflection and its nearly smooth posterior
slope.

There is a rather wide range of variation in the ratio of the length to the height of the
valves in the species, the length of the shell ranging from two and one half to nearly
three and one half times the height. Older specimens are usually higher and have much
heavier umbonal reflections.

Through the kindness of Dr. H. A. Pilsbry we were able to study and figure the type
of Pholas dilecta Pilsbry and Lowe. The holotype specimen appears to be only a rather
young, beach-worn specimen of P. chiloensis. Specimens from Kino Bay, Sonora, Mex-
ico labelled P. dilecta by Lowe are definitely this species.

Very little is known concerning this species beyond the fact that it has a rather wide

Plate 27. Pholas chiloensis Molina. Fig. 1. Pholas laqueata Sowerby (= P. chiloensis Molina) from west Colom-
bia, Hugh Cuming collector, Cotype (13x). Fig. 2. Kino Bay, Sonora, Mexico. H. N. Lowe collector
(natural size).

54 JOHNSONIA, No. 33 Zirfaea

range of distribution, extending as it does from northern Mexico to Chiloe Island, Chile.
Hupé stated that it was a rather common species on Chiloe Island, boring into soft stone.
Captain Phillip King who was on the HMS Adventure and the Beagle when those ships
were surveying the coast of South America stated ‘‘the soft parts of Pholas chiloensis are
considered very delicate by the inhabitants of the Island of Chiloe, by whom the animal
is called ‘Co-mes.” They are found in great abundance at low water imbedded in the
rocks near Sandy Point at San Carlos de Chiloe.”*’ Despite this apparent abundance in
restricted localities the species is generally rare in collections. We have seen only one
preserved specimen and most of the material studied was beach worn.

Range. From Kino Bay, Sonora, Mexico south to Chiloe Island, Chile.

Specimens examined. Mexico: Kino Bay, Sonora (MCZ; ANSP); Laguna de Scam-
mon, Baja California (ANSP); Santo Domingo, Baja California (USNM): Mazatlan,
Sinaloa: Acapulco, Guerrero (both MCZ). Nicaracua: Corinto (ANSP; W. J. Eyer-
dam). Panama: Panama City (MCZ; Univ. of Michigan); Pena Prieta, Panama City
(USNM). Ecuapor: between Manglaralto and Monte (USNM); Guayaquil (MCZ;
Redpath Museum). Peru: North of Payta(USNM); Payta(USNM; ANSP); Paraca
Bay (MCZ). Cute: Chiloe Island (AMNH).

Genus Zirfaea Gray

Zirfaea Gray 1840 [in] Synopsis of the Contents of the British Museum, ed. 42, p. 154 [nude name].

Zirfaea Gray 1842 [in] Synopsis of the Contents of the British Museum, ed. 44, p. 76 [defined, but no spe-
cies listed }.

Thurlosia ‘Leach’ Catlow and Reeve 1845, The Conchologist’s Nomenclator, p. 3 (genotype, Pholas crispata
Linné, monotypic).'

Zirfaea ‘Leach’ Gray 1847, Proceedings Zoological Society London, p. 188 (genotype, Pholas crispata
Linné, monotypic).

Zirphaea Leach 1852, A Synopsis of the Mollusca of Great Britain, p. 252 (genotype, Pholas crispata Linné,

monotypic).

Shell oval in outline, beaked anteriorly, rounded to truncate posteriorly, widely gap-
ing at both ends and having a sulcus extending from the umbo to the ventral margin.
There is a single accessory dorsal plate, the mesoplax, which is small and more or less
triangular in outline. Apophyses solid, strongly curved, broadened and often spoon-
shaped at the free end.

The genus Zirfaea is limited in its distribution to the colder waters of the Northern
Hemisphere. Zirfaea crispata Linné is known from both sides of the Atlantic, while Z.
pusbryi Stearns is restricted, so far as known, to the Pacific coast of North America.

Genotype, Pholas crispata Linné, subsequent designation, Gray 1847.

‘It is unfortunate that the name Thurlosia was introduced by Catlow and Reeve just two years before Gray
set the type for Zirfaea.. Should descriptions of genera with no species listed be invalidated in the future,
the name Thur/osia Catlow and Reeve 1845 will replace Zirfaea Gray 1842, unless this later name be placed
on the list of Nomina Conservanda.

Western Atlantic JOHNSONIA, No. 33

Ore
Gr

Zirfaea crispata Linné
Plates 1, 3 and 28 to 30

Mya crispata Linné 1758, Systema Naturae, ed. 10, 1, p. 670 (O. septentrionali),

Pholas crispata Linné 1776, Systema Naturae, ed. 12, 1, pt. 2, p. 111.

Pholas bifrons da Costa 1778, Historia Naturalis T'estaceorum Britanniae, p. 242, pl. 16, fig. 4 (Cornwall,
Lincolnshire, Yorkshire, Wales, etc. [British Isles] ).

Solen crispus Gmelin 1790, Systema Naturae, ed. 13, 1, pt. 6, p. 3228 (Teers [Tees River] Angliae).

Pholas parvus ‘Pennant’ Donovan 1800, The Natural History of British Shells 2, pl. 69 (non Barnea parva
Pennant).

Pholas crispa de Blainville 1825, Manuel de Malacologie, p. 578, pl. 79, fig. 7.

Zirfaea crispata Linné, Gray 1847, Proceedings Zoological Society London, p. 188.

Zirfaea crispata var. truncata Kaas 1939, Basteria 4, no. 1, p. 7, pl. 1 (Texel Island, Netherlands).

Distinctive characters. Shells white, beaked anteriorly, rounded to truncate posteriorly,
gaping at both ends and having an umbonal-ventral sulcus. Mesoplax very small and
triangular in outline. Valves usually in contact for only a short distance near the sulcus.
Siphons smooth, without the minute chitinous spots found in the Eastern Pacific species.

Description. Shells reaching 938.2 mm. (about 3? inches) in length and 49.2 mm. (about
2 inches) in height, beaked anteriorly, broadly rounded to truncate posteriorly, gaping
at both ends and with an umbonal-ventral sulcus. Valves usually in contact only at the
umbos and a small area on the ventral margin at the sulcus. In young specimens the
umbonal-ventral sulcus is quite pronounced and is evident on the inside of the valve as a
ridge. In older specimens the internal ridge is no longer visible, and the sulcus exists only

Plate 28. Zirfuea crispata Linné. Figs. 1-6. Rye Harbor, New Hampshire. Fig. 1. Shows the extent to
which the animal can withdraw into the shell. Fig. 2. Side view to show the expanded foot. Figs. 3-4.
Ventral views showing the way in which the animal can form various openings between the mantle and
the foot, through which streams of water can be directed into the anterior end of the burrow (all
natural size).

56 JOHNSONIA, No. 33 Zirfaea

as a Shallow furrow on the outside of the valve. In many young specimens this internal
ridge is swollen at the ventral margin forming an incipient condyle. Color a dull chalky-
white. Sculpture consisting of numerous concentric ridges which are laminated and strong-
ly imbricated anterior to the sulcus but are evident only as growth ridges on the posterior
slope. Radial ridges, expressed only by the radial arrangement on the rows of imbrica-
tions, are strong on the beaks, but are reduced in strength toward the umbos and posteri-
orly toward the suleus. In older specimens these imbrications may be reduced to undu-
lating ridges. Umbonal reflections broad and closely appressed over the umbos. Interior
of shells white and lightly glazed in young specimens while in older specimens it is usu-
ally chalky. Muscle sears and pallial sinus well marked, particularly in older specimens.
Pallial sinus broad and deep, and extending anteriorly about three fourths the distance
to the umbos. Apophyses solid, strongly curved, broadening at the free edge and becom-
ing quite spoon-shaped in some specimens. Protoplax lacking. Mesoplax small, fragile,
triangular in outline, and with a small basal flange. Periostracum thin, light straw-yellow
and deciduous.

The siphons are united and may be extended six to eight times the length of the shell.
The anterior one half to two thirds of the siphons are covered witha sheath of light straw-
yellow to medium-brown periostracum. ‘The remainder of the siphons lack the periostra-
cum and are white in color. The posterior ends of the siphons are marked with very fine
reticulations of dark mahogany-red. When the animal is extended, the reticulations are
widely spaced giving this region a gray-pink color. Incurrent siphon with dark red, pap-
illose and ciliated ridges running well into the siphonal tube. These ridges terminate in
branched cirri at the aperture. Excurrent siphon devoid of the internal ridges.

length height ratio h:]

93.5 mm. 49.2mm., 1.9 Small Point Beach, Maine

81.5 47.0 Naif Nahant, Massachusetts

70.0 42.5 1.6 Vaagnas Cape, Tromsé, Norway
45.0 28.0 1.5 Dover, England

34.0 14.0 2.4 Scituate, Massachusetts

30.0 23.0 1.3 Portland, Maine

29.5 27:0 1.4 Paratype of Z. c. var. truncata

Types, According to Hanley (1855, Ipsa Linnaei Conchylia, p. 26), the type of Mya
crispata Linné is in the Linnean collection. The type locality given by Linné was ‘‘O.
septentrionali.”’ It is here restricted to the banks of the Tees River, Middlesbrough,
England. This is the locality given by Lister 1678, the only publication to which Linné
referred for this species. The location of the types of Pholas bifrons da Costa is unknown.
The holotype of Zirfaea crispata var. truncata Kaas is in the collection of Dr. J. Th.
Henrard of Oegstgeest, Netherlands. Paratypes are in the collection of Dr. Piet Kaas
and the Museum of Comparative ZoGdlogy, no. 193521. The type locality is Isle of Texel,
Netherlands. We are grateful to Dr. Kaas for sending us a paratype specimen.

Remarks. Zirfaea crispata Linné is close in its relationship to Z. pilsbryi Lowe of the
Eastern Pacific. However, it may be distinguished from that species by its smaller size,
lighter, more fragile shell, less inflated posterior region, and by its smooth siphons which
are devoid of the minute chitinous spots usually found in Eastern Pacific species. In ad-
dition, the valves of Z. crispata Linné are usually in contact for only a short distance on

Western Atlantic JOHNSONIA, No. 33 ait

the ventral margin near the umbonal-ventral sulcus. The young specimens of these two
species are difficult and often impossible to distinguish. See remarks under Z. pilsbryt
Lowe.

The shells of Z. crispata Linné are variable in size and weight depending upon the con-
ditions under which the animals are living. Specimens living in soft, easily worked salt
marsh peat are elongate-oval in outline, and have rather widely spaced imbricated ridges
and fragile shells, indicating rapid growth. Those boring into rock or hard wood are
stunted, truncate, and have closely packed, badly worn imbricated ridges and heavy
shells, indicating hard work on the substratum and slow growth. It was such specimens
that were named Z. crispata truncata Kaas. Though this species is usually found in salt
marsh peat deposits, specimens have been found in a wide variety of material ranging
from stiff mud and blue clay to waterlogged wood, hard pine, poor quality bricks and
red sandstone. They are probably most abundant in the lower intertidal zone, but have
been taken alive in depths as great as 40 fathoms. They can withstand exposure to tem-
peratures ranging from below freezing to 26.6° C. However, as they live in the lower
intertidal zone or below the low tide level they are never exposed to the extremes of tem-
perature for an extended period of time. In laboratory experiments they can withstand
a rather wide range of salinity, but they are limited in nature to a marine habitat. Their
distribution is spotty though they may be exceedingly abundant in any one locality where
proper conditions exist. In one bed we counted over fifty specimens in a square foot of
peat. Many of the specimens were stenomorphs as their burrows were too close to allow
complete growth. The burrows were vertical and many extended toa depth of six inches
or more.

Zirfaea crispata Linné seldom bores into wood unless it is completely waterlogged and
softened. However, in a test block submerged at Beverly, Massachusetts, we found 56
minute specimens of this species. Among the thousands of test blocks that have been
examined from New England, this is only the second record for Zirfaea in the tests at
the Clapp Laboratories.

Little is known of the life history of this species, but according to Dr. C. M. Sullivan
(1948) the larvae occur in the plankton at Malpeque Bay, Prince Edward Island, from
the middle of June to the middle of July, and at that time the temperature of the water
ranges from 15° to 22° C. See also the introduction for further notes and figures of the
young.

Range. Easrern Aruantic: from Tromsé, Norway south to Boulogne, France.
WesteERN AtrLANtIC: from Labrador south to Anglesea, New Jersey. The record given
by C. W. Johnson, 1934 for South Carolina appears to be in error.

Specimens examined. FKasvERN ATLANTIC: Norway: Vaagnas Cape, Tromsé Sound
(Trondheim Museum); Tromsé (W. J. Eyerdam); Lofoten (USNM). Bririsu Istes:
Oban (MCZ); South Rock, St. Andrews, Fife (Gatty Marine Laboratory); Liverpool
(MCZ: AMNH); Hilbre Point, Dee River (Charleston Museum); Dover; Scarborough
(both MCZ). Neruertanps: Isle of Vlieland (A. N.C. ten Broek); Isle of Texel; Was-
senaar, about 6 miles north of The Hague (both P. Kaas); Scheveningen, The Hague
(D. Smits). Betcium: Nieuport (W. J. Eyerdam). France: Boulogne (MCZ).

58 JOHNSONIA, No. 33 Zirfaea

WeEsTERN ATLANTIC: LaBrapor: (Redpath Museum). NEwrounDLAND: (USNM).
QueBeEc: Gaspé (Redpath Museum); Coffin Island, Magdalen Islands (MCZ). Prixcr
Epwarpb IsLanp: Malpeque Bay (J.C. Medcof); Governor’s Island (Redpath Museum).
CarE Breron Istanp: (USNM). Nova Scorra: Straits of Northumberland (G. H.
Cox); Halifax (Redpath Museum); Digby (J. Bradley); off Digby in 40 fathoms (J.
Magarvey); Sable Island (USNM). Marine: Small Point Beach (A. Clarke); Penobscot
Bay (MCZ); Casco Bay (MCZ; Charleston Museum); Schoodic Point, Winter Harbor
(G. Moore); Portland: Kennebunk Beach (both MCZ); Skillings River, Lamoine (J.
Rankin): Jewels Island, Casco Bay; Isle of Shoals; John’s Bay, Bristol; Broad Sound
(all USNM). New Hampsuire: Rye Harbor (G. Moore). Massacnuserrs: Plum Is-
land, Newburyport: Ipswich; Salem; Beverly; Lynn; Swampscott; Nahant (all MCZ);
Salisbury (A. Clarke); Chelsea( AMNH); Winthrop; Scituate; Manomet Point; North
Beach, Orleans; Monomoy, Chatham (all MCZ); Hull; Duxbury (both MCZ; USNM).
RuopeE Istanp: Newport (ANSP). New York: Sag Harbor, Long Island (USNM);
Noyack Bay, Long Island (R. Latham). New Jersey: Sandy Hook (AMNH); As-
bury Park; Point Pleasant, Atlantic City; Ocean City; Anglesea (all ANSP).

Plate 29. Zirfaea crispata Linné. Fig. 1. Paratype of Zirfaea crispata var. truncata Kaas (= Zirfaea crispala
Linné). Isle of Texel, Netherlands (14x). Fig. 2. Nahant, Massachusetts. An old specimen to show pal-
lial line and muscle scars (apophysis broken off) (natural size). Figs. 3-4. Rivermere, Scituate, Massa-
chusetts. Fig. 4. A young specimen to show the internal umbonal-ventral ridge which more or less dis-
appears in the adult (13x).

Zirfaea pilsbryi Lowe
Plates 30 to 34
Zirfaea gabbi of authors, not of Tryon.
Zirfaea pilsbryi Lowe 1981, Nautilus 45, p. 53, pl. 3, figs. 1-2 (Bolinas, California).
Distinctive characters. Shell beaked anteriorly, broadly rounded to truncate posteriorly,
gaping at both ends, and having a sulcus extending from the umbo to the ventral mar-

Western Atlantic JOHNSONIA, No. 33 59

gin. Mesoplax small and more or less triangular in outline. Valves usually in contact on
the ventral margin for most of their length posterior to the sulcus. Siphons usually with
minute chitinous spots.

Description. Shell reaching 122 mm. (about 4? inches) in length and 68 mm. (about
24+ inches) in height, beaked anteriorly, broadly rounded to truncate posteriorly and
widely gaping at both ends. Valves in contact with each other at the umbos and for most
of the ventral margin posterior to the pedal gape. Sulcus extending from the umbo ob-
liquely to the ventral margin. In young specimens this sulcus may be barely visible ex-
ternally but it is clearly indicated internally as a strong beaded rib. In adult specimens
the furrow is always distinct externally but may be indicated on the inside of the valve
only as a row of isolated beads or it may be completely lacking. Umbonal reflection
broad and appressed against the umbo. At the point of contact of the two valves the re-
flection is always worn away, even in young specimens, as this area acts as an incipient
condyle on which the valves rotate. Color a dull chalky-white to light-salmon. Sculpture
consisting of numerous concentric ridges which are laminated and strongly imbricated on
the anterior slope but are indicated only as growth lines posterior to the furrow. The
radial ribs are restricted to the anterior slope and are indicated only by the radial arrange-
ment of the imbrications which are produced where the concentric ridges and radial ribs
cross. In young specimens the imbrications are very strong on the ventral margin of the
anterior slope, reducing in strength toward the umbo and posteriorly toward the furrow.
In old though living specimens, the sculpture on the anterior slope may be reduced to
undulating ridges, the free edges of which are recurved, giving an appearance of thick-

Plate 30. Mesoplax of Zirfaea. Figs. 1-3. Zirfuea crispata Linné, Rye Harbor, New Hampshire. Fig. 1.
Dorsal view. Fig. 2. Ventral view. Fig. 3. Side view showing the basal flange which is generally not vis-
ible in either of the other views. Figs. 4-6. Zirfaea pilsbryi Lowe, Anaheim Bay, California, an old speci-
men. Fig. 4. Dorsal view showing the proportionately small basal flange typical of old specimens. Fig. 5.
Ventral view. Fig. 6. Side view showing the marked curvature of the basal flange. Figs. 7-9. Zirfaea pils-
bryi Lowe, Playa del Rey, California, a young specimen with a proportionately large basal flange typical
of young and fast growing specimens.

60 JOHNSONIA, No. 33 Zirfaea

ness. Interior of shell white to light-salmon in color and glazed. Muscle scars and pallial
sinus well marked. Pallial sinus broad and deep extending inward nearly to the umbo.
Apophyses rather large, solid, strongly curved, moderately to strongly spooned at the
ventral margin and marked with concentric growth lines. Protoplax lacking. Mesoplax
transverse, small but strong, more or less triangular in outline and usually with a well
developed basal flange. Periostracum thin, straw-yellow to red-brown in color, and usu-
ally persistent on the posterior slope.

Plate 31. Zirfuea pilsbryi Lowe, Alamitos Bay, California (about 2 natural size). Showing the minute chitinous
spots on the siphons and the extent to which the valves are in contact on the ventral margin.
(Photographs received through the kindness of J. E. Fitch, California Fisheries Laboratory. )

The siphons are united and they may be extended six to eight times the length of the
shell. The anterior one third of the siphons is covered with a dark-horn to red-brown
periostracum which also extends over the posterior end of the shell. ‘The remainder of the
siphons lacking periostracum, a gray-white to light-ivory in color, and usually irregularly
marked with small orange chitinous spots. These spots vary from one half to two milli-
meters in greatest diameter and never reach the size or thickness of those found on the
siphons of ovoidea Gould. The area immediately surrounding the siphonal openings is
marked with fine reticulations of dark mahogany-red. Incurrent siphon with dark-red,
branched cirri at the opening, and these are continued as ridges which extend a short dis-
tance within. Excurrent siphon with several minute ridges and small cirri. Foot nearly
circular in outline and truncate. Foot and mantle light-ivory in color. The above descrip-
tion of the soft parts is based upon preserved material.

length height ratio h:]

192°0%m. 67/.0'mm. 1.8 Newport, Oregon

115.5 Saree | Anaheim Bay, California

105.0 68.0 15 South Alki Beach, Seattle, Washington
99.5 51.0 1.9 Lisabeula, Washington
76.0 37.0 2.0 Holotype. Bolinas, California
61.5 84.5 1.8 San Diego, California

Types. The holotype of Zirfaea pilsbryi Lowe is in the Academy of Natural Sciences,
Philadelphia, no. 50809, from Bolinas, California, H. Hemphill, collector. Paratypes are
in the Lowe Collection which is now at the San Diego Museum of Natural History.

EE ——————————

Western Atlantic JOHNSONIA, No. 33 61

Remarks. Zirfaea pilsbryi Lowe is closely related to Z. crispata, but may be distin-
guished from that species by its larger size and by having the ventral margin of the
valves, especially in older specimens, nearly straight posterior to the pedal gape. This
allows the valves to come in contact for a considerable distance along the ventral margin
as shown in the figure of the living specimen. In Z. crispata the ventral margin is usually
in contact for only avery short distance near the base of the umbonal-ventral suleus. The
post-umbonal area of Z. pilsbryi is generally longer, more truncate and inflated than in
Z. crispata Linné, and in addition, the siphons are usually marked with small chitinous
spots.

A lot of Zirfaea received from Humboldt Bay, California, lacks the small chitinous
dises on the siphons, and in other respects these specimens more closely resembled Z.
crispata of the Western Atlantic. Ina letter, J. EK. Fitch of the California Fisheries Lab-
oratory informed us that the east coast oyster has been imported at this locality and he
stated that there are now in the bay large beds of Mya and Venus and a number of gas-
tropods all of which came in with the seed oysters of Crassostrea. Consequently it may
well be that both species of Zifaea now occur at this locality.

Zirfaea pilsbryt Lowe was formerly known as Z. gabbi Tryon. Lowe examined the
type of gabbi and found that it was not in the genus Zirfaea, but was the young stage of
a Penitella, so. he placed it in the synonymy of Penitella penita Conrad. For a further
discussion of this species see the remarks under Penitella penita Conrad and P. gabbi
Tryon which will be covered in Part II of this family.

Zirfaea pilsbryi Lowe is usually found boring into mud and clay banks, the larger
specimens burrowing to a depth of ten to fourteen inches (Fitch 19538, p. 95). Occasion-
ally, like its Western Atlantic counterpart, it is found boring into decaying, waterlogged
wood.’ Specimens from the Queen Charlotte Islands which had been boring into sand-

Plate 32. Zirfaea pilsbryi Lowe, from Bolinas, California. Holotype (slightly enlarged).

'W.K. Emerson 1951, Bulletin Southern California Academy of Sciences 50, pt. 2, p. 89-91.

62 JOHNSONIA, No. 33 Zirfaea

stone were stunted and misshapen. An interesting account of the ecology and method
of boring of this species is given by MacGinitie (1935, pp. 731-735).

Range. From Nunivak Island, Alaska south to San Jaunico Bay, Baja California
(Fitch 1953, p. 95).

Plate 33. Zirfaea pilsbryi Lowe. Fig. 1. Bolinas, California. To show muscle scars and pallial line, umbonal-
ventral ridge barely visible. Fig. 2. A truncate form from San Diego, California (both about # natural size).

Specimens examined, ALASKA: Nunivak Island; Port Méller; Bering Island (all
USNM); Cordova (W. J. Eyerdam). Brirish CoLumMBia: Queen Charlotte Islands
(USNM). Wasuincron: Port Orchard; South Alki Beach, Seattle; Anacortes; Brain-
bridge, near Winslow, Kitsap County (all W.J. Eyerdam); Fort Lawton Beach, Puget
Sound; Lisabeula (both MCZ); Willapa Bay (EK. P. Chace). Orrecon: Netarts Bay,
Tillamook County (R. Coats); Newport (ANSP). Canirornia: Daby Island and Buhne
Point, Humboldt Bay (both J. E. Fitch); Bolinas(MCZ; ANSP; USNM); San Fran-
cisco; Santa Cruz (both MCZ); Monterey Bay (EK. P. Chace); Elkhorn Slough, Mon-
terey Bay (J. E. Fitch); San Pedro Bay (AMNH); Anaheim Bay (E. P. Chace;
USNM; MCZ); San Diego (AMNH; USNM); Playa del Ray (W. J. Eyerdam);
Alamitos Bay, Long Beach (J. E. Fitch). Mexico: Bahia San Quintin and Punta
Abreojos, Baja California [dead and worn specimens] (both USNM).

Plate 34. Zirfaea pilsbryi Lowe, Marin County, California. Ventral view of opposed valves to show the
apophyses and hinge area (about 14x).

Western Atlantic JOHNSONIA, No. 33 63

Notes

Barnea subtruncata Sowerby

Two additional Mexican records for this species were received from Dr. S. S. Berry
after the section on this species had been printed. These are San Felipe and Punta
Cyote, northwest of La Paz, both Baja California in the Gulf of California.

Barnea maritima ‘d° Orligny> Dall

Barnea maritima ‘d*Orbigny’ Dall 1889, Bulletin United States National Museum, no. 37, p. 72 (Texas)
[nomen nudum].

We have been unable to find a description of this species by d’Orbigny and _ believe
that it was possibly a manuscript name on specimens in the collection of the United
States National Museum which Dall had used at the time he worked up his list of the
marine mollusks of the southeastern states. Since that time this name has appeared in
several lists.

Book Reviews

Abbott, R. T. 1954: American Sea Shells. D. Van Nostrand Co., Inc., New York,
pp. 14+541, 40 plates (24 in color), 100 text figures. Abbott has presented the many
students and collectors of our American marine mollusks with a very readable and in-
formative book. He considers the shells of our east coast from Labrador to Texas, includ-
ing only those West Indian elements that reach Florida, and the west coast from Alaska
to southern California. For the professional worker it presents a source of information
and answers many queries which arise almost daily. To the beginner and the ‘‘professional
amateur’’ it will be helpful in determining their local material as well as giving sugges-
tions on procedure in caring for the collection. Few, if any, of our common species have
been overlooked and many of the rarer species have been included. The colored plates
are excellent and the black and white plates are, for the most part, exceedingly clear as to
detail. Many of the text figures are by J. C. McConnell who was for many years asso-
ciated with Dall. Few artists have had his ability in pen drawing.

There is much in addition to the taxonomy and descriptive elements in this book.
Abbott has included in detail, methods of collecting, life histories, habitats and many
other subjects not presented in popular books on mollusks. Chapter 14 is a guide to the
literature which is most valuable for locating additional information about our marine
mollusks.

The scientific names of our mollusks change with the acquisition of new knowledge.
Many new names or new name combinations appear in this book with no explanation for
these various changes. This is unfortunate. These must either be accepted blindly or
considered individually and restudied, a long and thankless task and a great duplication of
effort. However, this criticism is trivial, considering the book as a whole. It is very well
done and will do much to advance knowledge among an ever increasing number of com-
petent students.—W. J. CLENCH

64 JOHNSONIA, No. 33 Book Reviews

La Rocque, Auréle 1953, Catalogue of the Recent Mollusca of Canada. National
Museum of Canada, Ottawa, Bulletin no. 129, 9+406. This is an exceedingly important
piece of research. The catalogue proper is arranged taxonomically from Classes down to
Subgenera. Each species has the original citation and one or more references to standard
North American monographs or other studies covering the species in question. Type
localities are given as well as the known range for each species. The ranges for the ma-
rine mollusks of both coasts are now pretty well known, as well as for most species of
land shells. The fresh-water species, however, are far less known and many of the ranges
given by La Rocque are open to question. This is not at all the fault of La Rocque, but
it is because the available literature on our fresh-water mollusks is mainly old, particularly
papers of monographic scope, and many of the published data are quite unreliable. The
author has a peculiar and somewhat misleading procedure which is to cite a species under
its present genus followed by the original reference without the original name combina-
tion. For example:

**Paravitrea multidentata Binney 1840, Journal Boston Soc. Nat. Hist., vol. 3, p. 425.7"

The genus Paravitrea was established by Pilsbry in 1898, a name, of course, unknown
to Binney in 1840. Such entries are readily recognized in the present paper as La Rocque
has dated the establishment of each genus. ‘The danger here is that subsequent authors,
not having a large library to consult, are very apt to quote as original from La Rocque
such name combinations which were not given in the original citation.

There is an excellent bibliography and a complete alphabetic index.—W. J. CLENcH

Carcelles, Alberto R. 19538, Catalogo de la Malacofauna Antarctica Argentina. Anales
del Museo Nahuel Huapi 3, pp. 150-250, 5 plates and 1 map. This is a companion re-
port to the three earlier papers on the mollusk fauna of Argentina by Carcelles which
were reported upon in Johnsonia 2, p. 880. This present report covers the Antarctic
fauna and lists 338 species included in 175 genera. As in previous reports, the author has
given a brief history of the work accomplished in this area. The catalogue is arranged
systematically, and for each species there are several references, the known range of the
species and the localities for the specimens which he examined. An excellent table is
given showing the geographic distribution of the species of this region and their occur-
rence as well in the Magellanic Province. The plates are excellent and 114 species have
been illustrated. This very important piece of work when combined with Carcelles’ three
other papers, gives a complete summation of the Argentinian marine mollusks. There is
also an extensive bibliography.—R. D. TuRNER

ti

JOHNSONIA

Published by
THe DEPARTMENT oF MOLLUSKS

Museum of Comparative Zodlogy, Harvard University

Cambridge, Massachusetts

MARCH 29, 1955 PHOLADIDAL VOL. 3, NO. 34

THE FAMILY PHOLADIDAE
IN THE WESTERN ATLANTIC AND THE EASTERN PACIFIC

PART II—MARTESIINAE, JOUANNETIINAE AND XYLOPHAGINAE
BY
Rutu D. Turner’

In the introduction to Part I of this study there is a general discussion of the classifi-
cation of the Family Pholadidae and the various subfamilies of which it is composed.
Also included in the first part are general data regarding life histories, ecology, distribu-
tion, methods of boring, anatomy, physiology and economic importance.

Since the publication of Part I, Dr. N. T. Mattox of the Allan Hancock Foundation
has kindly sent all of their material for study. Most of the records have been incorporated

Plate 35. Damage caused by Martesia striata Linné and Teredinidae. Section of a test board which was sub-
merged at San Juan, Puerto Rico on Sept. 21, 1944 and removed on March 21, 1945. The long tubes are
the work of teredo or shipworms. Views of the anterior ends of specimens of Martesia striata Linné with
the callum fully developed show the funnel-shaped pits formed by the umbonal reflections. Note that Mar-
tesia does not avoid the teredo tubes (14x).

'The family Pholadidae, Parts I-II, were submitted in partial fulfillment of the requirements for the de-

gree of Doctor of Philosophy, Dept. of Biology, Harvard University.

65

66 JOHNSONIA, No. 34 Chaceia

in the main portion of this paper. Records for the species covered in Part I will be in-
cluded in a report for the Allan Hancock Foundation. A key to the genera of the Pho-
ladidae is included at the end of the paper.

Subfamily MARTESIINAE
Genus Chaceia, new genus’

Shells moderate to large in size, gaping widely at both ends in the young stage and
only partially closing the pedal gape with a callum in the adult stage. Shell divided into
two parts by a pronounced umbonal-ventral sulcus. Valves, when in normal position,
usually in contact for only a short distance on the ventral margin near the base of the
umbonal-ventral sulcus. Protoplax lacking, being replaced by a dorsal extension of the
callum. Mesoplax small, transverse, in one piece and similar to that found in Zzrfaea.
Metaplax and hypoplax lacking. Siphons large, incapable of retraction within the shell.
Foot in young specimens broadly oval in outline and truncate, atrophied in the adult.

The genus Chaceia differs from other closely related genera in this subfamily by having
the shell gaping at both ends, by lacking a siphonoplax and by having only a partial callum.

There is only one known species in this genus and this is limited in its distribution to
California and Mexico.

Genotype, Pholas ovoidea Gould

Chaceia ovoidea Gould
Plates 36-39

Pholas ovoidea Gould 1851, Proceedings Boston Society Natural History 4, p. 87; Gould 1853, Boston
Journal of Natural History 6, p. 388, pl. 15, fig. 1 (Monterey, California).

Pholadidea ovoidea Gould, Gould and Carpenter 1856 [1857] Proceedings Zoological Society London, p. 198.

Parapholas ovoidea Gould, H. and A. Adams 1856, Genera of Recent Mollusca 2, p. 330.

Pholadidea ovoidea Gould, Oldroyd 1924, Stanford University Publications, Geological Sciences 1, no. 1,
p. 212, pl. 21, figs. 5-6 (not pl. 51, fig. la-b); Fitch, J.E., 1953, State of California, Department of Fish

and Game, Fish Bulletin no. 90, p. 93, fig. 59.

Distinctive characters. Shell gaping widely at both ends in young specimens and only
partially closing the pedal gape with a callum in the adult. Growing edge of the callum
infolded over the beaks. Mesoplax similar to that found in Zirfaea. Shell divided into

Plate 36. Chaceia ovoidea Gould. White Point, San Pedro, California (natural size). Young working speci-
mens. Fig. 1. External view of valves showing the pronounced constriction of the shell at the umbonal-
ventral suleus. Fig. 2. Internal view of valve showing the sulcus expressed internally as a ridge. Fig. 3.
Ventral view of opposed valves showing the pedal gape extended to its fullest extent.

an : - - : = ( June ;
Named in honor of Mr. E. P. Chace of Lomita, California.

Eastern Pacific JOHNSONIA, No. 34 67

two portions by avery pronounced umbonal-ventral sulcus, the anterior portion tapering
to a point on the ventral margin at the sulcus. Siphons very large, not capable of retrac-
tion within the shell, having large oval to elongate orange chitinous patches, and being
strongly papillose to warty at the tip.

Description. Shell reaching 115.5 mm. (about 44 inches) in length and 71 mm. (about
2? inches) in height, broadly oval in outline, inflated, rather light in structure but strong
and producing a partial callum in the adult stage. Immature specimens strongly beaked
anteriorly with a nearly circular pedal gape. Posteriorly they are broadly rounded and
widely gaping. Shell divided into two well defined areas by a pronounced umbonal-
ventral sulcus. Anterior portion sculptured with close-set, upturned, undulating, concen-
tric ridges and rather weak radial ribs which are indicated mainly by the radial arrange-
ment of the rows of undulations. Young specimens occasionally have imbrications pro-
duced where the concentric ridges and radial ribs cross one another. Posterior portion
sculptured with strongly marked concentric ridges which are definite extensions through
the sulcus of the ridges of the anterior slope. Umbonal reflections simple, rather broad,
closely appressed over the umbos, but usually free anteriorly. In adult specimens the
pedal gape is partially closed by the callum which is sculptured by fine growth lines. The
callum extends dorsally between the beaks and over the umbonal reflections and, doub-
ling upon itself, forms a partial enclosure for the anterior adductor muscle. Between the
beaks the callum is folded inward. Protoplax absent, being replaced by a dorsal exten-
sion of the callum. Mesoplax rather small, transverse, broadly V-shaped, in one piece and

Plate 37. Chaceia ovoidea Gould. Anaheim Bay, California (natural size). Upper figure, inside of valve of an
adult specimen showing the infolding of the callum over the beaks, the muscle scars and the pallial sinus.
Lower figure, outside of valve showing strong umbonal-ventral sulcus and pronounced ridges on the posterior
slope.

JMAR 29 yore

RARYERD

RM vrnerey

JOHNSONIA, No. 34 Chaceia

~
oO
D

somewhat similar to that found in Zrfaea. Occasionally in old specimens the dorsal mar-
gin of the mesoplax appears to be extended anteriorly as a result of the deposition of cal-
cium in the periostracum covering the area between the mesoplax and the callum. In-
terior of the shell white and usually glazed. Umbonal-ventral sulcus evident internally
as a pronounced ridge. Muscle scars well marked, especially in older specimens. Pallial
sinus extending anteriorly beyond the umbonal-ventral ridge. Apophyses small, solid,
strong, and extending from beneath the umbos at an angle paralleling the umbonal-
ventral ridge. Periostracum thin, light straw-yellow in color and persistent.

Siphons very large, not capable of retraction within the shell, and they may be ex-
tended 6 to8 times the length of the shell. Anterior portion of the siphons covered with
a moderately heavy golden to red-brown periostracum which extends dorsally up between
the valves and ventrally covers that area in the adult not covered by the callum. Pos-

ify

%,

7

Sez

tN

Plate 38. Chaceia ovoidea Gould. Figs. 1-2, White Point, San Pedro, California. Fig. 1. Specimen showing
the beginning of the production of the callum and the siphons contracted to their fullest extent. Fig. 2.
Ventral view of a young specimen showing valves in normal position, coming in contact only at the base of
the umbonal-ventral ridge, the point on which the valves rock when boring (about 14x). Fig. 3. Adult
specimen with fully developed callum from Carpenteria, California (about 2 natural size). Note the large
chitinous patches and the warty appearance of the tip of the siphons.

All photographs received through the kindness of J. E. Fitch of the California Fisheries Laboratory.

Eastern Pacific JOHNSONIA, No. 34 69

terior to the periostracal covering, the siphons are grayish-white and have pronounced
oval to long, crescent-shaped, orange-brown chitinous patches. These patches are some-
what irregularly spaced but are most abundant near the posterior end. The tip of the
siphon is dark red-brown to dark purple in color and strongly papillose to warty. The
siphonal apertures are circled with white. Foot in young specimens large, broadly oval
in outline and truncate. It is atrophied in the adult. Foot and surrounding mantle white
to light purple-gray in color.

length * height ratio h:]

11525) mm). 71.0 mm. 1.62 Carpenteria, California

103.0 70.0 1.47 7
85.0 45.0 1.88 Anaheim Landing, California
faeo 48.0 LoS Bahia San Bartolomé, Baja California
55.5 38.0 1.46 ene mf He cl

* All specimens measured were adults

Types. ‘The location of the type specimen of Pholas ovoidea Gould is unknown. The
type locality is Monterey Bay, California, Major Rich, collector.

Remarks. Chaceia ovoidea Gould is a very distinctive and easily recognized species.
When the entire animal is at hand, the very large siphons with conspicuous orange chit-
inous patches and the incomplete, infolded callum easily distinguish it from all other spe-
cies. The young stage is most easily confused with young, truncate specimens of Zirfaea
pulsbryi Lowe, the shells of the two species having about the same weight and the perios-
tracum of both being thin and light straw-yellow in color. In C. ovoidea Gould, however,
the two portions of the shell are much more clearly marked, the umbonal-ventral sulcus
being much more pronounced, resulting in a marked constriction of the shell. In addi-
tion, the anterior portion of C. ovoidea Gould tapers to a point on the ventral margin at
the sulcus. The sculpture of the posterior portion of C. ovoidea Gould is stronger than
that of Z. pilsbryi Lowe and the posterior slope is far more inflated. The sculpture on
the anterior slope of Z. pilsbryi Lowe consists of rather strong, pointed imbrications
whereas in C. ovoidea Gould the sculpture consists of upturned, undulating ridges. From
young of Penitella penita Conrad and Penitella gabbi Tryon this species differs by having
a much more inflated posterior slope, a much thinner periostracum, a wider pedal gape

Plate 39. Mesoplax of Chaceia ovoidea Gould. Figs. 1-3. Dorsal, ventral and side view of the mesoplax of an
adult specimen from Bahia San Bartolomé, Baja California. This shows the great similarity of the dorsal
plate in Chaceia to that in Zirfaea. Figs. 4-6. Dorsal, ventral and side view of the mesoplax from a young
specimen taken at White Point, San Pedro, California.

TO JOHNSONIA, No. 34 Penitella

and in having the apophyses paralleling the umbonal-ventral ridge rather than projecting
anteriorly from beneath the umbos at a sharp angle. From the adults of these two species
it differs by having an incomplete callum. From Penitella fitchi Turner, the only other
species in the eastern Pacific with an incomplete callum, C. ovoidea differs by its siphons,
its mesoplax and by lacking the siphonoplax.

There has been considerable confusion as to the identity of this species, probably due
to the fact that C. ovoidea Gould isa rather rare species with a somewhat restricted range
and because Gould’s figure and description are rather poor. Unless one has an adult
specimen of ovoidea, the figures of Gould could easily be interpreted as either Penitella
penita Conrad, without its siphonoplax, or as P. gabbi Tryon. The matter was further
confused when I. Oldroyd (1924, pl. 51, figs. la-b) figured P. gabbi Tryon as ovoidea
Gould.

Mr. E. P. Chace has written that at White Point, San Pedro, California, C. ovoidea
Gould lives in soft shale rock along with P. penita Conrad, P. gabbi Tryon and Para-
pholas californica Conrad, but specimens are much less abundant. He collected some
large specimens with Mr. J. Fitch at Carpenteria, California and found that they made
an excellent chowder. According to Mr. Fitch, ovordea may bore to a depth of 18 to 21
inches, which is much deeper than that of the species with which it is found living.

Range. From Santa Cruz, California (J. E. Fitch, 1958, p. 93) south to San Diego,
California and at Bahia San Bartolomé, Baja California.

Specimens examined. CALIFORNIA: Santa Barbara (MCZ); Carpenteria (E. P. Chace;
J. KE. Fitch: A. Hancock Foundation); White Point, San Pedro (KE. P. Chace); Ana-
heim Landing (W. J. Eyerdam); San Diego (ANSP; MCZ). Mexico: Bahia San
Bartolomé, Baja California (J. E. Fitch).

Genus Penitella Valenciennes

Penitella Valenciennes 1846 [in] Abel du Petit-Thouars, Voyage Autour du Monde sur la Frégate La Vénus,
Atlas de Zoologie, Mollusques, plate 24.

Penicilla ‘Valenciennes’ Conrad 1854, Journal Academy Natural Sciences Philadelphia (2) 2, p. 335 (error
for Penitella Valenciennes).

Shells small to moderate in size, the larger species reaching about 95 mm. (about 3?
inches) in length, oval in outline, divided into two distinct regions by an umbonal-ventral
suleus and producing a callum in the adult stage. Shell beaked and widely gaping an-
teriorly in the young stage, with a nearly circular pedal gape. Valves rounded to truncate
and closed posteriorly, the siphon being capable of complete retraction within the shell.
Umbonal reflections variable, ranging from those which are thin and very closely ap-
pressed so that the sculpture of the shell shows through, to those which are heavy and
free anterior to the umbos. Protoplax lacking, being replaced by the dorsal extension of
the callum. Mesoplax transverse, in one piece and, in young specimens, very similar to
that found in Zirfaea. In the adult, a dorsal portion is added to the mesoplax which
grows forward and encloses the posterior portion of the anterior adductor muscle. Metaplax
and hypoplax lacking, Siphonoplax variable, present or absent. Siphonal tube lacking.

Penitella Valenciennes has been considered a subgenus of Pholadidea by many authors.
However, the form of the mesoplax, the type of siphonoplax, the complete lack of a

Eastern Pacific JOHNSONIA, No. 34 Ta

siphonal tube and the absence of any indication of a metaplax or hypoplax seem sufficient
reason to consider this group of species as a genus. In Pholadidea, the mesoplax is
divided longitudinally into two parts and the two halves of the siphonoplax are somewhat
fused to form a tube. In Penitella the mesoplax is in one piece and the two portions of
the siphonoplax are never fused and are often diverging. In addition, the apophyses of
the Pholadidea are very small, narrow and fragile, while those of Penitella, though often
short, are heavy, expanded and blade-like at the free end.

Unfortunately Stoliczka in 1870, Dall in 1898, and Lamy in 1926, were in error when
they considered P. penita Conrad as the type of the genus Penitella. This species was
not included by Valenciennes at the time the genus was instituted by him.

The genus Penitella Valenciennes, so far as now known, is restricted in its distribution
to the Eastern and Northern Pacific.

Genotype, Penitella conradi Valenciennes, subsequent designation Habe 1952.

Key to the species of Penitella

(based on adult specimens)

i. Aqdults-with a complete callam . . .. . 2 2. ee. te Se 2
Adults with.an incomplete callam. . . . 2. 40.6 5% ie _ fitehi

2. Umbonal reflection closely appressed for the entire length, et onan
present, siphons smooth. . ... . oes ee sf Sd hs eae

Umbonal reflection free anteriorly, a een nnias Sreaie. coe pustu-

LOSE MGS sins os ae : Pa Tee asc > hy OE
3. Siphonoplax Se of — flexible ere nutie® flaps which are not
lined with caleareous granules. Mesoplax pointed posteriorly, truncate
anteriorly and having lateral wings . . . ie feo ea, 5 emai
Siphonoplax not flexible, composed of a hese faeces outer layer lined
with coarse calcareous granules. Mesoplax truncate posteriorly, pointed
anteriorly and lacking lateral wings . . . ....... . . . . conradi

Penitella fitchi, new species'
Plates 40-42

Distinctive characters. Shell having thin, very closely appressed umbonal reflections,
producing only a partial callum in the adult stage and having the siphonoplax composed
of numerous chitinous leaf-like layers. Mesoplax broadly rounded to truncate posteriorly,
rounded anteriorly and lacking lateral wings.

Description. Shell white, oval in outline, reaching 49.5 mm. (about 2 inches) in length
and 35.5 mm. (about 12 inches) in height, solid in structure and producing a partial cal-
lum and a siphonoplax in the adult stage. Immature specimens beaked and gaping widely
anteriorly, rounded and closed posteriorly. Valves divided into two distinct and very un-
equal parts by a pronounced umbonal-ventral sulcus. Anterior portion small, about 4 the
posterior portion, tapering to a point on the ventral margin at the umbonal-ventral wlan
and sculptured by very close-set, upturned undulating ridges and radial ribs. The ribs are
expressed mainly by the radial arrangement of the undulations. Posterior portion broadly
rounded in young specimens, becoming proportionately longer and more tapering in the

This species is named for John E. Fitch of the California Fisheries Laboratory, Terminal Island Station,
who has been most helpful in obtaining material for us and who collected this species.

72 JOHNSONIA, No. 34 Penitella

adult and sculptured with fine growth lines. Umbos located near the anterior third of the
shell and covered, in young specimens, with broad, simple umbonal reflections which are
closely appressed for their entire length and through which the sculpture of the shell can
be seen. In the adult, the pedal gape is partially closed by a rather heavy, solid callum
which extends dorsally up between the beaks and over the umbonal reflection to form the
enclosure for the anterior portion of the anterior adductor muscle. The callum is sculp-
tured with growth lines and there is a slight indication of the forward extension of the
radial ribs. Anteriorly a broad oval area remains after the callum is completed and this
is covered with a heavy gray-brown periostracum, only a minute pore actually remaining
open in the adult stage. Protoplax lacking, being replaced by the dorsal extension of the
callum. Mesoplax in young specimens broad, thin, nearly flat, rounded posteriorly and
with a deep embayment anteriorly. In the adult stage the mesoplax extends dorsally and
anteriorly forming an enclosure for the posterior portion of the anterior adductor muscle.
Mesoplax of the adult large, heavy, more or less triangular in outline, rounded _posteri-
orly and pointed anteriorly. Siphonoplax composed of numerous leaf-like layers of gray-
brown chitin, the leaves becoming progressively longer from the inside out. These ex-
tend well up the dorsal margins of the valves nearly to the umbos, and on the ventral
margin extend nearly to the umbonal-ventral sulcus. The number of layers increases with
age, a young specimen Just beginning to produce a callum usually having only two layers
while an old mature specimen may have eleven or more. Metaplax and hypoplax lacking.

Plate 40. Penitella fitchi Turner. Bahia San Bartolomé, Baja California. Fig. 1. Paratype. Ventral view to
show the partial callum. Fig. 2. Holotype. Dorsal view to show the triangular mesoplax and the multi-
layered siphonoplax. (Both about 2x. )

Eastern Pacific JOHNSONIA, No. 34 73

Periostracum thin, medium gray-brown to yellow-brown in color, and deciduous. In-
terior of shell white and glazed. Umbonal-ventral sulcus visible internally as a broad,
smooth ridge. Muscle scars large, pronounced and roughened. Pallial sinus broad and
deep, extending anteriorly beyond the umbonal-ventral ridge. Apophyses solid, expanded
and blade-like at the free end and extending anteriorly from beneath the umbos at a
sharp angle.

Siphons united, white to ivory in color, smooth and capable of complete retraction
within the shell. Incurrent siphon with six large and numerous small cirri which extend
for a short distance internally as ridges. Excurrent siphon about one half the diameter
of the incurrent siphon and with numerous small cirri. Externally the tips of the siphons
are marked with a narrow band of dark red-brown vermiculations and spots. Foot and
mantle white to ivory. In young specimens the foot is large, nearly circular in outline
and truncate. It is atrophied in the adult. The above description of the soft parts is based
upon preserved material.

length * height ratio h:]
49.5 mm. 35.5 mm. 1.4 Holotype
40.0 35.0 ial Paratype
38.0 21.5 1.7 oy
16.5 11.5 1.4 e7.

* All specimens measured had a callum

Plate 41. Penitella fitchi Turner. Bahia San Bartolomé, Baja California. Fig. 1. Lateral view to show the
small anterior portion and the siphonoplax. Fig. 2. Internal view of adult showing the incurving of the
callum over the beaks, the muscle scars and the apophysis. Fig. 3. Internal view of young specimen show-
ing the proportionately much shorter posterior slope, the muscle scars and the expanded apophysis. Fig.
4. External view of young specimen showing the very closely appressed umbonal reflection and the triangu-
lar shape of the anterior portion. Fig. 5. Ventral view of opposed valves showing the small ventral condyle,
the apophyses and the pedal gape. (All paratypes, all natural size. )

74 JOHNSONIA, No. 34 Penitella

Types. Holotype, Museum of Comparative Zodlogy, no. 189413, from Bahia San
Bartolomé, Baja California, John E. Fitch, collector. Paratypes from the same locality
collected by J. E. Fitch, E. Dwyer and D. Joseph, are in the Museum of Comparative
Zoblogy, the United States National Museum, Stanford University, the Academy of
Natural Sciences Philadelphia and the collection of Dr. S. S. Berry.

Remarks. Penitella fitehi Turner is a very distinctive species, especially in the adult
stage, and it does not appear to be closely related to any other species in this genus. It
is, perhaps, most closely related to Penitella conradi Valenciennes as both species have a
very similar young stage, and the mesoplax in the young and adult stage of both species
are close in general outline. However, P. fitchi Turner differs from P. conradi Valen-
ciennes by having only a partial callum and by having the siphonoplax composed of
numerous thin chitinous leaves. Penitella penita Conrad, a species which P. fitchi Turner
approaches in size and with which it is found living, differs in both the form of the meso-
plax and the siphonoplax and in having a complete rather than partial callum. From
Chaceia ovoidea Gould, the only other species which produces an incomplete callum, P.
fitchi Turner differs by having a siphonoplax and by having the shell closed posteriorly,
the siphons being capable of complete retraction within the shell. In addition, the siphons
of P. fitchi Turner are smooth while those of C. ovordea Gould have large, orange, chit-
inous patches.

Mr. John E. Fitch has written that P. fitchi Turner was boring into the sedimentary
rocks on the north side of Bahia San Bartolomé [Turtle Bay], Baja California along with
equal numbers of P. penita Conrad and a few C. ovoidea Gould. They appeared to bore
to about the same depth as P. penita. The outstanding difference between the two spe-
cies seen in situ is the greater amount of color on the siphons of penita—the siphons of
jitchi appearing creamy white when extended.

Range. Known only from the type locality, Bahia San Bartolomé, Baja California.

4mm.

Plate 42. Mesoplax of Penitella fitchi Turner. Fig. 1. Dorsal view of the mesoplax showing the truncate pos-
terior margin and the pointed anterior end. Fig. 2. Ventral view of the mesoplax showing the large cavity.
Fig. 3. Side view of the same specimen. Fig. 4. Dorsal view of the mesoplax of a young specimen. All
from specimens collected at Bahia San Bartolomé, Baja California.

Eastern Pacific JOHNSONIA, No. 34 75

Specimens examined. Mexico: Bahia San Bartolomé, Baja California (J. EK. Fitch).

Penitella conradi Valenciennes
Plates 48-46; Plate 72, figs. 1-2

Penitella conradi Valenciennes 1846 {in| Abel du Petit-Thouars, Voyage Autour du Monde sur la Frégate
La Vénus, Atlas de Zoologie, Mollusques, plate 24, fig. 1 (no locality given); Lamy 1921, Bulletin Muséum
d’Histoire Naturelle Paris 27, p. 179 (Monterey, California, from type label).

Navea subglobosa Gray 1851, Annals Magazine Natural History (2) 8, p. 385 (California, in hole in a shell);
H.and A. Adams 1856, The Genera of Recent Mollusca 2, p. 328, pl. 89, fig. 6a-b; Lamy 1925, Journal de
Conchyliologie 69, p. 103.

Martesia intercalata Carpenter 1855, Catalogue of the Reigen Collection of Mazatlan Mollusea, p. 13 (Mazat-
lan, Mexico in Spondylus lamarckii).

Navea newcombii Tryon 1865, American Journal of Conchology 1, p. 39, 285, pl. 2, figs. 1-3 (Lower Cali-
fornia in Haliolis, W. Newcomb, collector).

Penitella parva Tryon 1865, American Journal of Conchology 1, p. 39, pl. 2, figs. 4-5 (Lower California,
in Haliotis, W. Newcomb, collector).

Distinctive characters. Shell usually found boring into Halotis or other shells; occa-
sional specimens are found in clay and soft stone. Umbonal reflections broad and closely
appressed for their entire length. Mesoplax truncate posteriorly, pointed anteriorly and
lacking lateral wings. Siphonoplax heavy, not diverging and composed of a chitinous
outer layer which is lined with a white, granular, calcareous deposit.

Plate 43. Penitella conradi Valenciennes. Fig. 1. A copy of the original figures of the species as given by
Valenciennes in the ‘Voyage de la Vénus,’ pl. 24, fig. 1. Fig. 2. Side view of a type specimen. Fig. 3.
Dorsal view of the type.

Photographs of the type specimens were received through the kindness of Dr. Max de Jardin of the Mu-
seum d’Histoire Naturelle, Paris, France.

76 JOHNSONIA, No. 34 Penitella

Description. Shell reaching about 88 mm. (14 inches) in length and 16.5 mm. (about
+ inch) in height, generally oval in outline, solid in structure and producing a callum and
siphonoplax in the adult stage. Immature specimens beaked and widely gaping anteriorly,
tapering posteriorly to a rounded and closed posterior margin. Shell divided into two
distinct regions by a pronounced umbonal-ventral sulcus. Anterior portion about one
third the posterior portion, tapering to a point on the ventral margin at the umbonal-
ventral sulcus, and sculptured with very fine, close-set, upturned, undulating, concentric
ridges and weak radial ribs which are expressed mainly by the radial arrangement of the
undulations. Posterior portion sculptured with concentric growth lines. Umbonal re-
flections wide, closely appressed and with thickened, elevated, longitudinal ridges over
the umbos. In adult specimens the pedal gape is closed by a heavy callum which is sculp-
tured with very faint growth lines and faint ridges indicating the extension forward of
the radial ribs. Protoplax lacking, being replaced by the dorsal extension of the callum.
Mesoplax in young specimens thin, nearly flat, and more or less semicircular in outline.
In adult specimens the dorsal portion of the mesoplax extends anteriorly and encloses
the posterior portion of the anterior adductor muscle. It is broadly rounded to truncate
posteriorly and pointed anteriorly, fitting in between the two halves of the reflected cal-
lum. Metaplax and hypoplax lacking. Siphonoplax dark brown in color, covered with

Plate 44. Penitella conradi Valenciennes. Figs. 1-3. Del Monte Beach, Monterey Bay, California. Specimens
boring in soft shale (about 3x). Figs. 4-8. Monterey Bay, California. Specimens boring in Haliotis rufes-
cens brought in at Monterey. Such specimens are always small and more or less deformed (about 2x). Figs.
2 and 8 show the white lining of the siphonoplax. Apophysis broken in fig. 2. Figs. 3 and 6 show the typi-
cal mesoplax which is pointed anteriorly and lacks lateral wings.

Eastern Pacific JOHNSONIA, No. 34 ar

a heavy periostracum and lined internally with a white, granular, calcareous deposit. The
two halves of the siphonoplax come together to form a cup-like structure which is very
variable both in length and diameter. Periostracum especially heavy on the posterior
slope, dark brown to nearly black in color and persistent. Interior of shell white and
generally chalky. Umbonal-ventral sulcus evident internally as a rather weak rib. Muscle
sears large, pronounced and usually roughened. Pallial sinus broad and deep, extending
anteriorly beyond the umbonal-ventral ridge. Apophyses solid, expanded and blade-like
at the free end, and extending from beneath the umbos anteriorly at a sharp angle, often
nearly reaching the anterior margin of the shell. Occasionally the apophyses are rather
deformed and irregular.

Siphons united, small, white and devoid of periostracum. They probably do not ex-
tend more than half the length of the shell and are capable of complete retraction within
the shell. Aperture of the incurrent siphon surrounded with numerous cirri which are
flecked with dark red-brown markings. Excurrent siphon about one half the diameter of
the incurrent siphon, having much smaller cirri and fewer, smaller flecks. Foot and man-
tle white to light ivory in color. Foot in young specimens large, nearly circular in out-
line and truncate; it is atrophied in the adult. The above description of the soft parts
was based upon preserved material.

length* height ratio h:]
33.0 mm. 16.5 mm. 2.0 Del Monte Beach, Monterey, California (in shale)
30.0 16.2 8 in - ‘" - pe chs
17.8 es jes From Haliotis rufescens landed at Monterey, California
521 8.8 1.8 it * v raoto # 5
8.1 5.5 1.5 = gs 563 Ne wae ne “3
6.8 4.5 1.5 a s * Tes a ee

* All specimens measured had a complete callum

Plate 45. Penitella conradi Valenciennes. Figs. 1-2. Lectotype, Penitella parva Tryon (=P. conradi Valen-
ciennes) Baja California (about 44x). Figs. 3-4. Holotype, Navea newcombi Tryon (=P. conradi Valencien-
nes) Baja California (about 4 $x). Figs. 5-9. Monterey Bay, California from Haliotis (about 24x). Fig. 5.
Specimen with partially grown callum and showing the beginning of the elongation of the posterior slope.
Fig. 6. Young specimen. Fig. 7. Internal view of young specimen showing the large muscle scars, pallial
sinus and apophysis. Fig. 8. View of pedal gape to show the blade-like apophyses projecting forward.
Fig. 9. Ventral view of opposed valves showing the condyles and closed posterior margin.

78 JOHNSONIA, No. 34 Penitella

Types. The types of Penitella conradi Valenciennes are in the Paris Museum; the type
locality is Monterey, California, from the type label (Lamy 1921). The types of Navea
subglobosa Gray are in the British Museum; the type locality is California. The lectotype
of Penitella parva Tryon is in the Academy of Natural Sciences Philadelphia, no. 50999,
from Lower California, obtained from a Haliotis, W. Newcomb, collector. The holotype
of Navea newcombu Tryon from the same locality is also in Philadelphia, no. 51069. The
location of the types of Martesia intercalata Carpenter is unknown. They are not in the
Hanley collection in the British Museum. However, a sketch made by P. P. Carpenter
of the specimens in the Hanley collection is in the British Museum and we reproduce a
photograph of this sketch. Specimens so named by Carpenter are in the United States
National Museum and the Redpath Museum, Ontario, Canada.

Remarks. Penitella conradi Valenciennes is close in its relationship to P. penita Conrad
and P. gabbi Tryon, and has been confused with both these species. It is the species com-
monly found in Hadliotis, but may also be found boring into other shells or into soft rock
or clay along with P. penita Conrad and P. gabbi Tryon. From adult P. penita Conrad
it differs by having a heavier, less diverging siphonoplax with a pronounced white, gran-
ular, calcareous lining. The mesoplax of P. conradi is truncate posteriorly and pointed
anteriorly, fitting in between the reflected portions of the dorsal extension of the callum,
and in addition, lacks the lateral wings found in the other two species. From P. gabbi
Tryon it differs by having very closely appressed umbonal reflections and in having a
siphonoplax. The young of P. conradi Valenciennes are rather difficult to distinguish from
the young of other species in this genus. From the young of P. gabbi Tryon it is distin-
guished by its umbonal reflection and from the young of P. penita Conrad, which also
has a wide and closely appressed umbonal reflection, it can be distinguished by the apophy-
ses, which generally project forward at a much sharper angle, by the elevated longitudinal
ridges over the umbos, the shorter, more rounded, posterior slope and the more upturned
concentric ridges on the anterior portion. Young specimens of P. conradi Valenciennes
boring into clay and soft rock are most difficult to distinguish, but the shorter, more
rounded, posterior slope and the shape of the mesoplax will separate them.

Plate 46. Mesoplax of Penitella conradi Valenciennes. Fig. 1. Dorsal, ventral and side view of the mesoplax
of a specimen taken from shale, showing the pointed anterior end. Fig. 2. Dorsal, ventral and side views
of the mesoplax of a much malformed specimen taken from a Haliotis. Figs. 3-4. Plates from more typical
specimens boring in Haliotis. The series is given to show the range of variation that may be expected.
Fig. 5. Dorsal view of the mesoplax of a young specimen taken from a Haliotis. Fig. 6. Ventral view of
the same specimen. All specimens are from Monterey Bay, California.

Eastern Pacific JOHNSONIA, No. 34 79

ie

Two problems have confused the nomenclature of this species: one, the generally ac-
cepted idea that P. parva Tryon (=P. conradi Valenciennes) was restricted to boring
into shells, and the other, the lack of understanding of the young form of this species.
Probably as a result of the idea that the ‘‘Haliotis borer’’ was never found in clay and
rock, Lamy (1921) considered P. conradi Valenciennes as a synonym of P. penita Con-
rad. Though Valenciennes never described Penitella conradi, his figures clearly show the
characteristic mesoplax which is broadly rounded to truncate posteriorly, pointed anteri-
orly and lacks lateral wings. ‘The umbonal reflections and the heavy, cupped, rather than
diverging siphonoplax are also characteristic of the ‘‘Ha/liotis borer.”* In the large series
of specimens, mostly P. penita Conrad, which were taken from the rocks at Del Monte
Beach, Monterey Bay, California and Bahia San Bartolomé, Baja California we have
found specimens of the ‘‘Haliotis borer’’ which match perfectly the illustrations by
Valenciennes, On the basis of the material now available, there seems to be no question
that P. parva Tryon and P. conradi Valenciennes refer to the same species—Tryon’s ma-
terial having come from a Hadliotis while Valenciennes’ specimens came from soft shale.
Specimens boring into shells are nearly always deformed, often with the posterior portion
and the siphonoplax greatly reduced. The specimens may be curved and dented to fit
around obstructions in their burrows and the anterior portion of the shell is often deformed
and flattened. In heavily infected Haliotis the specimens often run into each other’s bur-
row. Regardless of whether the specimens are malformed ones from Hadliotis or perfect
ones from a clay bed, the characteristic mesoplax and siphonoplax as well as the granular
muscle scars are always evident.

The young or working form of P. conradi Valenciennes has never been figured, and
though specimens of this species with a half grown callum are rare we have seen several
and figure one of them. By removing the callum and siphonoplax from numerous adult
specimens of P. conradi Valenciennes we were able to show that the young stage of this
species is identical with what has generally been referred to as Navea subglobosa Gray.
It is interesting to note the P. parva Tryon and Navea newcomli Tryon were both taken
from Haliotis collected in Baja California by W. Newcomb. Very possibly they both
came from the same shell. This problem of placing the young stages of callum-building
pholads in separate genera has appeared several times and undoubtedly before the entire
family is completely studied several more species and genera will fall into synonymy.

Penitella conradi Valenciennes has been found boring into Haliotis fulgens Philippi,
H. rufescens Sowerby, and H. sorenseni Bartsch, Mytilus californianus Conrad, Astraea
sp., Pododesmus sp. and in soft rock. Mr. John Fitch has written that the borers do rela-
tively little damage to the Haliotis so far as the food crop is concerned though they can
be very abundant in some specimens.

Range. From Gualala, Mendocino County, California south at least as far as Bahia
San Bartolomé, Baja California.

Specimens examined. CALIFORNIA: Gualala, Mendocino County (USNM): Stewarts
Point and Shell Beach, Sonoma County (Hancock Foundation): Farallones, off San
Francisco (Redpath Museum; USNM); Monterey Bay (MCZ; USNM: E. P. Chace:
W.J. Eyerdam); Del Monte Beach, Monterey, in shale (EK. P. Chace): Pacific Grove
(S. S. Berry); Point Sur, Monterey County in 10 fathoms (M. Keen); 5 miles north of

80 JOHNSONIA, No. 34 Penitella

Morro Rock (J. E. Fitch); White Point, San Pedro (J. E. Fitch; E. P. Chace; S. S.
Berry: USNM); Laguna Beach, Orange County (S. S. Berry); 15 mi. east of Cardwell
Point, San Miguel Island; Anacapa Passage, Santa Cruz Island in 50 fathoms (both
Hancock Foundation); Santa Catalina Island (USNM); San Clemente Island (J. E.
Fitch): off San Clemente Island in 10 fathoms (S. S. Berry). Mexico: off South Coro-
nado Island: Isla Geronimo; Punta Santo Tomas and Santo Tomas Anchorage, all Baja
California (all S. S. Berry): Bahia San Bartolomé, Baja California (J. E. Fitch; S.S.
Berry): Todos Santos Bay, Baja California (ANSP).

Penitella penita Conrad
Plates 5, 47-51

Pholas penita Conrad 1837, Journal Academy Natural Sciences Philadelphia 7, p. 237, pl. 18, fig. 7 (San
Diego and Santa Barbara, California).

Pholas concamerata Deshayes 1839, Revue Zoologique par la Société Cuviérienne, p. 357; Deshayes 1840,
Magasin de Zoologie, Guérin-Méneville (2) 2, pl. 17.

Penitella spelaea Conrad 1855, United States Pacific Railroad Exploration and Survey, California. Prelimi-
nary Geological Report 8, Appendix, pp. 319 and 326, pl. 5, fig. 43 a—b (San Pedro, California, Recent for-
mation).

Penitella penita Conrad, Tryon 1862, Proceedings Academy Natural Sciences Philadelphia 14, p. 215.

Pholadidea penita Conrad, Carpenter 1864, Report British Association Advancement of Science for 1863, p.525.

Penitella curvata Tryon 1865, American Journal of Conchology 1, p. 40, pl. 2, figs. 6-8 (Straits of Fuca
[Vancouver], W. M. Gabb, collector).

Pholadidea sagitta Stearns’ Dall 1916, Proceedings United States National Museum 52, p. 417 (Monterey,

California) [young specimen |.

Distinctive characters. Callum protruding beyond the beaks. Umbonal reflections
broad, thin and very closely appressed their entire length. Mesoplax sharply pointed

Plate 47. Penitella penita Conrad. Figs. 1-2. Anaheim Bay, California. Adult specimen with fully developed
callum, siphonoplax missing. Fig. 2. Internal view showing muscle scars and pallial sinus. Fig. 3. Netarts
Bay, Tillamook County, Oregon. A small, but adult specimen with fully developed callum and siphono-
plax. Figs. 4-5. Monterey Beach, Monterey, California. Young working specimens. Fig. 4. Showing the
very closely appressed umbonal reflection. Fig. 5. Ventral view of opposed valves showing the pedal gape
and the ventral condyle. (All about natural size. )

Eastern Pacific JOHNSONIA, No. 34 81

posteriorly, truncate anteriorly and having short, pointed, lateral wings. Siphonoplax com-
posed of two heavy, flexible, chitinous flaps which are often diverging. Siphons smooth.

Description. Shell medium to rather large in size, reaching 93 mm. (about 3? inches)
in length and 49 mm. (about 2 inches) in height, oval in outline, solid in structure and
producing a callum in the adult stage. Immature specimens beaked and gaping widely
anteriorly, tapering slightly posteriorly to a broadly rounded posterior margin which, in
adult forms, becomes truncate and terminates in a chitinous flap or siphonoplax. Shell
divided into two distinct regions by an umbonal-ventral sulcus. Anterior beaked portion
sculptured by close-set, upturned, undulating, concentric ridges and indistinct radial ribs
which are indicated largely by the radial arrangement of the undulations. Posterior por-
tion sculptured only by close-set growth lines. Umbos prominent, located near the an-
terior third of the shell and covered by broad, thin umbonal reflections which are closely
appressed their entire length, through which the sculpture of the shell surface can be
seen. In adult specimens the pedal gape is closed by a rather heavy callum which ex-
tends slightly beyond the beaks and which is sculptured with weak concentric growth
lines and faint parallel ridges. A narrow slit remains when the callum is completed
and this is covered by a heavy periostracum, with only a minute pore remaining open
anteriorly. The callum is extended dorsally over the umbonal reflection and, doubling
upon itself, encloses the anterior adductor muscle. Protoplax lacking, being replaced by
the dorsal extension of the callum. Mesoplax in young specimens triangular and deeply
V-shaped. In adult specimens the dorsal portion of the mesoplax extends anteriorly to
form the enclosure for the posterior portion of the anterior adductor muscle. It is usu-
ally truncate anteriorly, pointed posteriorly, sharply keeled ventrally and has two small,
pointed, lateral wings. Siphonoplax consisting of leathery chitinous flaps which are often
widely diverging; they may vary in color from a light yellowish-brown to dark horn color
and range in length from 6 to 15 mm. on specimens 65 to 70 mm. long. There are no
other accessory plates. Periostracum, particularly on the posterior slope, rather heavy
and ranging from a light brown to almost black in color.

Interior of shell white and usually glazed. Umbonal-ventral sulcus evident internally
as a low ridge which, in young specimens, extends slightly beyond the ventral margin
of the shell forming a condyle. Muscle scars well marked, pallial sinus broad and deep
extending inward anterior to the umbonal-ventral ridge. Apophyses rather short, solid,
generally flattened at the free end and projecting from beneath the umbos anteriorly at
a moderate to sharp angle.

Plate 48. Penitella penita Conrad. Pholas curvata Tryon (=P. penita Conrad) from Straits of Juan de Fuca,
Washington. Lectotype (54x).

89 JOHNSONIA, No. 34 Penitella

The siphons may be extended about the length of the shell, and they are capable of com-
plete retraction within the shell. Siphons smooth, white for nearly the entire length, but
usually marked with a narrow band of dark red-brown reticulations at the posterior ex-
tremity. Incurrent siphon with six large and several small branched cirri surrounding the
aperture and extending inward as ridges for a short distance. Excurrent siphon with a
few very small cirri and few or no internal markings. Foot and mantle white. Foot in
immature specimens large, oval in outline and truncate. In adult specimens the foot
atrophies and the mantle is closed except for a minute pore at the anterior extremity.
The description of the soft parts is based upon preserved material.

The following series of measurements show the variation in the proportion of the length
to the height of the shell. All specimens measured had a fully developed callum. The
siphonoplax was not included in the measurement of the length as it is so variable and

is often broken.

Plate 49. Penitella penita Conrad. White Point, San Pedro, California. Figs. 1 and 3. Adult specimens with
fully developed callum and siphonoplax. Fig. 2. A young working specimen showing the large, nearly
circular foot which atrophies in the adult stage (about 1.8x).

Photographs received through the kindness of J. E. Fitch, California Fisheries Laboratory.

Eastern Pacific JOHNSONIA, No. 34 83

length * height ratio h:]
93.0 mm. 49.0 mm. 1.9 Port Orchard, Oregon
74.5 23.0 3.2 Buhne Point, Humboldt Bay, California
63.4 21.8 2.9 Netarts Bay, Tillamook, Oregon
48.0 27.0 1.8 Anaheim Bay, California
17.0 8.0 rl White Point, San Pedro, California
10.0 4.0 2.5 pie as a ee

* All specimens measured had a complete callum.

Types. The location of the type specimen of P. penita Conrad is unknown. The type
locality is San Diego, California. The lectotype of P. curvata Tryon is in the Academy
of Natural Sciences Philadelphia, no. 51004, from the Straits of Fuca [Vancouver],
W. M.Gabb, collector. The location of the type of Pholas concamerata Deshayes is un-
known; the type locality is California. The holotype of Pholadidea sagitta ‘Stearns’ Dall
from Monterey Bay, California is in the United States National Museum, no. 68312.

Remarks. This species is one of the most common and best known of the pholads in
the Eastern Pacific, In the adult stage it is readily identified by its flexible, leathery
siphonoplax, its characteristic mesoplax, and its smooth siphons. Young specimens may

Plate 50. Penitella penita Conrad. White Point, San Pedro, California. Fig. 1. Dorsal view of an adult show-
ing the mesoplax which is pointed posteriorly, and the broad umbonal reflection extending on either side
of the dorsal extension of the callum (about 24x). Fig. 2. Dorsal view of young showing the mesoplax.
Figs. 3-4. Holotype of Pholadidea sagitta Dall (=P. penita Conrad), Monterey Bay, California (a young
specimen about 2.3x).

84 JOHNSONIA, No. 34 Penitella

be confused with the young of P. gabbi Tryon and of C. ovoidea Gould, but can usually
be distinguished by the broad, closely appressed umbonal reflection. Specimens of P. con-
radi when boring into soft stone are quite similar to penita but the mesoplax of the for-
mer species lacks the lateral wings and the siphonoplax is not flexible. See remarks under
P. gabbi Tryon and P. conrad Valenciennes.

There has been considerable confusion as to the identity of P. sagitta ‘Stearns’ Dall,
as it was very briefly described and unfortunately was never figured. After examining
the type, which we figure, we believe it to be nothing more than a young specimen of
P. penita.

Penitella penita may be found in a variety of substrata ranging from stiff blue clay to
sandstone and cement. Specimens are very variable both in size and shape depending
upon the amount of crowding and the hardness of the substratum. Examples from Buhne
Point, Humboldt Bay, California and some from Netarts Bay, Tillamook, Oregon, are
unusually long in proportion to their height, probably showing evidence of rapid growth
in an easily worked substratum. Rapidly growing specimens usually have a thin shell
with a thin, light colored periostracum and a more fragile callum. Specimens boring into
harder rock may become badly distorted especially at the anterior end, and in addition,
the shells may be curved and misshapen to fit around obstructions in their burrow.
Tryon’s P. curvata was based on such specimens.

In the final report of the San Francisco Bay Marine Piling Investigation, Kofoid and
Miller (1927) stated that P. penita Conrad did considerable damage to concrete jacketed
piling in San Francisco Bay and Los Angeles Harbor. They also state that P. penita
Conrad is edible and may be used for food where it occurs in sufficient numbers to justify
getting them out of the rocks.

Range. From Bering Island, Siberia south at least as far as Bahia San Bartolomé,
Baja California.

Specimens examined: SiBeER1A: Bering Island (USNM). Atasxa: Chirikof Island
(USNM); Yakataga Reef, Yakataga District (Hancock Foundation). British CoLum-
BIA: Queen Charlotte Islands (USNM); Straits of Juan de Fuca, Vancouver (ANSP;
USNM); Victoria (ANSP); Nanaimo (MCZ). Wasuincton: Neeah Bay (USNM);
Port Orchard; South Alki Beach, Seattle (both W. J. Eyerdam); Willapa Bay (E. P.

Plate 51. Mesoplax of Penitella penita Conrad. Figs. 1-2. Dorsal and ventral views of the mesoplax of a young
specimen to show the relationship to Zirfaea. Fig. 3. Dorsal view of the mesoplax of an old, worn adult
specimen. Fig. 4. Ventral view of the mesoplax of an average adult specimen showing the cavity into which
the posterior end of the anterior adductor muscle fits. Fig. 5. Dorsal view of the same specimen to show
the dorsal portion which is well inset on the broad base of the young stage forming the lateral wings. Fig.
6. Side view of the mesoplax of an adult specimen. All from specimens collected at Tillamook, Oregon.

Eastern Pacific JOHNSONIA, No. 34 85

Chace; MCZ); Culvers Point, San Juan Island (EK. Swan). Orecon: Netarts Bay (R.
Coats); Yaquina Bay (MCZ); Newport (MCZ; ANSP; W.J.Eyerdam); Coos Head,
Coos Bay (MCZ; USNM): South and Middle Bays, Cape Arago (Hancock Founda-
tion). CALIFORNIA: Crescent City; Half Moon Bay (both MCZ); Pt. Arena; Carmet;
Shell Beach; Stewarts Point; Bodega Lagoon; Tomales Bluff (all Hancock Founda-
tion); Monterey Bay (MCZ; USNM; ANSP; Hancock Foundation); Del Monte
Beach, Monterey Bay (EK. P. Chace); off Del Monte in 12 fathoms (S. S. Berry); San
Luis Obispo; Santa Barbara (both MCZ); San Miguel Island (USNM): 14 miles east
of Cardwell Point, San Miguel Island (Hancock Foundation); Carpenteria; White
Point, San Pedro (both E. P. Chace and J. K. Fitch); San Pedro (USNM; MCZ; E.P.
Chace); Long Beach (MCZ); Buhne Point, Humboldt Bay (J. E. Fitch); Anaheim
Bay (MCZ; Hancock Foundation; W.J.Eyerdam); Hazard Canyon, 7 to 8 miles south
of Morro Bay (J. E. Fitch); Point Loma (USNM); Portuguese Bend (Hancock Founda-
tion); San Diego (MCZ; ANSP; USNM). Mexico: Colnett, Baja California (Han-
cock Foundation); Bahia San Bartolomé, Baja California (J. E. Fitch; USNM).

Penitella gabbi T?-yon
Plates 52-54
Zirfaea gabbi Tryon 1863, Proceedings Academy Natural Sciences Philadelphia, p. 10, pl. 1, fig. 1 (Coast
of Japan ?); H. N. Lowe 1931, Nautilus 45, p. 52.

Pholadidea ovoidea ‘Gould’ Oldroyd 1924, Stanford University Publications. Geological Sciences 1, no. 1,
pl. 51, fig. la—b, not the text or plate 21, figs. 5-6; non ovoidea Gould.

Distinctive characters. Callum not protruding beyond the beaks. Umbonal reflections
narrow, lightly appressed over the umbos and free anteriorly. Mesoplax in the adult
specimens with a rounded point posteriorly, pointed anteriorly and with broad lateral
wings. Siphonoplax lacking. Siphons pustulose.

Description. Shell oval in outline, reaching 55 mm. (2 inches) in length and 30 mm.
(1¢ inches) in height, solid in structure and producing a callum in the adult stage. Im-
mature specimens beaked anteriorly and gaping widely, rounded posteriorly and closed.
Shell divided into two distinct areas by an umbonal-ventral sulcus. Anterior portion
sculptured by close-set, upturned, undulating, concentric ridges and radial ribs. These
ribs are indicated largely by the radial arrangement of the undulations and are usually
arranged in pairs. Posterior portion sculptured only with concentric growth lines. Um-
bos prominent and usually located near the anterior third of the shell, however, the post-
umbonal length of the shell is rather variable depending upon the speed of growth and
the type of substratum in which the animal is boring. Umbonal reflections narrow, free
for most of their length and only lightly appressed over the umbos. In adult specimens
the pedal gape is closed by a callum which is sculptured with fine concentric growth lines
and usually with parallel ridges which indicate a forward extension of the radial ribs.
The callum extends dorsally between the beaks, but does not protrude anterior to them
or imbed them; it encloses the anterior portion of the anterior adductor muscle. Only
a very narrow slit remains when the callum is completed and this is covered by a heavy
periostracum with only a minute pore remaining open in the adult. Protoplax lacking,
being replaced by the extension of the callum over the umbonal area. Mesoplax trans-

86 JOHNSONIA, No. 34 Penitella

verse and in young specimens deeply V-shaped. In the adult a dorsal portion is added to
the mesoplax to enclose the posterior portion of the anterior adductor muscle. This upper
portion varies with age from truncate to pointed anteriorly. There are two wide lateral
wings on the adult mesoplax. Siphonoplax lacking. Periostracum moderately heavy, a
dull grayish-brown to medium brown in color and persistent.

Interior of shell white and usually glazed. Umbonal-ventral sulcus indicated internally
as a strong ridge which terminates ventrally in asmall condyle. Muscle scars well marked,
pallial sinus broad and deep and extending anteriorly to beyond the umbonal-ventral
ridge. Apophyses rather short, solid, enlarging slightly at the free end and extending
from beneath the umbos anteriorly at a rather sharp angle. Periostracum not extending
inside the shell at the siphonal area.

The siphons may be extended 1 to 2 times the length of the shell and may be con-
tracted completely within the shell. They are devoid of periostracum, white in color and
strongly pustulose. The pustules are irregular in size and distribution on the siphons and
there is no indication of chitinous caps covering the pustules as in P. ovordea Gould. On
the specimens studied the pustules appeared to be most abundant near the mid portion,
the area near the shell and a small band near the openings being nearly smooth. On occa-
sional specimens there may be faint reticulated markings of a dark mahogany brown at
the tip of the siphons. Surrounding the opening the incurrent siphon has 8 to 10 large
and several small cirri which extend inside as ridges. Excurrent siphon about one half the
diameter of the incurrent siphon, nearly smooth and devoid of all color internally. Man-
tle and foot white. Foot in young specimens nearly circular in outline and truncate. In
adult specimens the mantle is closed except for a minute pore anteriorly and the foot
atrophies. The description of the soft parts is based upon preserved material.

Plate 52. Penitella gabbi Tryon. Fig. 1. Restoration Point, Puget Sound, Washington. Figs. 2-3. Del Monte
Beach, Monterey Bay, California (all natural size). Fig. 2. Internal view showing the apophysis and muscle
sears. Figs. 1 and 3 show the range of length-height proportion, the rounded posterior margin, the cavity
beneath the umbonal reflection, and the grouping of the rows of imbrications. Figs. 4 and 5 are young
specimens showing the free umbonal reflection and the ventral condyle.

Eastern Pacific JOHNSONIA, No. 34 87

length* height ratio h:]
Do.o mim. 30 mm. 1.8 Holotype of gabbi Tryon
48.0 25 1.9 Del Monte Beach, Monterey Bay, California
32.0 18 1.8 Culvers Point, San Juan Island, Washington
22.0 12 1.8 4 mile south of White Point, San Pedro, California

* All specimens measured had a complete callum

Types. The holotype of Zirfaea gabbi Tryon [ = Penitella gabbi) is in the Academy of
Natural Sciences Philadelphia, no. 51085. In the original description the locality was
given as ‘Japan?’ but the label on the type specimen gives California, W. M. Gabb, col-
lector. The type locality is here restricted to Monterey Bay, California, an area from
which we have seen specimens and where Gabb had collected.

Remarks. Adult specimens of P. gabbi Tryon differ from P. penita Conrad, the species
with which it is most closely related, by lacking the siphonoplax, by having a more nar-
rowly rounded posterior margin, a shorter, broader mesoplax, and a lighter, thinner perios-
tracum. In addition, the umbonal reflections of P. gabbi Tryon are narrow, free anteriorly
and only lightly appressed over the umbos, while in P. penita they are broad, very closely
appressed and almost approach a glaze. In P. gabbi the callum is not built forward of the
beaks and does not imbed the beaks as it does in most species in this genus. Young
specimens of P. gabbi and P. penita are often difficult to distinguish but the narrow, free
umbonal reflections and the more tapering posterior slope of the former can usually be
depended upon to separate this species. If the soft parts can be studied the pustulose
siphons of gabbi quickly distinguish it from P. penita.

From Chaceia ovoidea Gould this species differs by having a heavier, less inflated
shell, a smaller pedal gape and a much more tapering, narrowly rounded posterior slope
which is not gaping. In addition, the apophyses of P. gabbi project from beneath the
umbos anteriorly at a rather sharp angle, whereas in C. ovoidea Gould they parallel the
umbonal-ventral ridge. The mesoplax in the two species are very different as shown in
the plates. In the adult stage Chaceia ovoidea Gould is usually much larger and the cal-
lum is not completely closed. If the soft parts are available, the large orange chitinous

Plate 53. Penitella gabbi Tryon. Fig. 1. Culvers Point, San Juan Island, Washington. Preserved specimen
showing the pustulose siphons, the broad wings on the mesoplax and the lack of the siphonoplax (about
12x). Fig. 2. Holotype of Zirfaea gabbi Tryon (= Penitella gabbi Tryon) (natural size),

88 JOHNSONIA, No. 34 Penitella

patches on the siphons of C. ovoidea Gould are in strong contrast to the white, pustulose
siphons of P. gabbi Tryon.

There has been considerable confusion concerning Penitella gabbi Tryon. The name
was based on a single mutilated valve from an uncertain locality. On examining the type
it was obvious that it was not a Zrfaea but the adult of a Penitella with the callum
broken away. Tryon, not realizing this was a callum-producing form, originally described
it as a Zirfaea. Lowe later synonymized it with P. penita Conrad and many other authors
have confused it with C. ovoidea Gould. Oldroyd (1924, pl. 51, fig. la—b) figures a good
specimen of P. gabbi Tryon showing the knobby pustulose siphons, but unfortunately
under the name of P. ovoidea Gould.

The young of P. gabbi Tryon are very close in appearance to Zirfaea constricta Sow-
erby, a Japanese species. When more material is available and the life history of Z. con-
stricta is known, it may prove to produce a callum when adult; it would then belong in
the genus Penitella in which case the two species may be the same.

Penitella gabbi Tryon is found in the same general habitat as P. penita Conrad, but is
in general, a much rarer species. This rarity is probably responsible for the apparent gaps
in its distribution.

Range. From Drier Bay, Alaska south to San Pedro, California.

Specimens examined. ALASKA: Drier Bay (W. J. Eyerdam). Brirish CoLuMmBIA:
Nanaimo (MCZ). WasnHineron: Tacoma(USNM); Restoration Point, Puget Sound;
Port Orchard (both W.J.Eyerdam); Culvers Point, San Juan Island (EK. Swan). Ore-
GON: Coos Bay (Hancock Foundation). Catirornia: Bodega Lagocn (Hancock Foun-
dation); Monterey Bay (MCZ): Del Monte Beach, Monterey Bay (E. P. Chace); off

Del Monte in 12 fathoms (S. S. Berry); White Point, San Pedro (J. E. Fitch): 4 mile
south of White Point (E. P. Chace); Santa Barbara (MCZ): Hazard Canyon, 7 to 8

miles south of Morro Bay (J. E. Fitch); San Diego (ANSP; Hancock Foundation).

Plate 54. Mesoplax of Penitella gabbi Tryon. Fig. 1. Dorsal view of the mesoplax of a young specimen. Fig.
2. Ventral view of the same specimen. Fig. 3. Side view of the same specimen. Fig. 4. Dorsal view of the
mesoplax from a specimen that was just beginning to produce a callum showing that at the same time the
dorsal portion of the mesoplax is formed. Fig. 5. Ventral view of the same specimen showing a curious de-
formation of the ventral ridge. Fig. 6. Dorsal view of the mesoplax of an adult specimen to show the
pointed anterior end. Fig. 7. Ventral view of the same specimen. Fig. 8. Side view of the same specimen.
All figures are from specimens taken at Monterey Bay, California.

Eastern Atlantic JOHNSONIA, No. 34 89

Genus Pholadidea Turton

Pholadidea ‘Goodall’ Turton 1819, A Conchological Dictionary of the British Islands, London, p. 147.

Pholadidoidea de Blainville 1826, Dictionnaire des Sciences Naturelles 39, p. 535 (genotype, P. goodall
Turton [ =loscombiana Turton], monotypic).

Cadmusia Leach 1852, A Synopsis of the Mollusca of Great Britain, p. 254 (genotype, Cadmusia solanderia
Leach [ =/oscombiana Turton] monotypic).

Shells small to moderate in size, more or less oval in outline, divided into two distinct
regions by an umbonal-ventral sulcus, and producing a callum in the adult stage. An-
teriorly the shell is beaked and widely gaping in the young stage, rounded and closed pos-
teriorly. Umbonal reflections variable. Protoplax lacking, being replaced by a dorsal
extension of the callum. Mesoplax small to moderate in size, divided longitudinally into
two parts and with the protruding basal portion present or absent. Incipient metaplax
and hypoplax present or absent, not existing as separate plates but resulting from the
deposition of calcium in the periostracum extending between the valves posterior to the
umbos. Siphonoplax largely chitinous and usually fused to form a tube. An additional
calcareous siphonal tube is present in some species. Apophyses very small, fragile and not
broadening at the free end. Siphons capable of complete retraction within the shell.

Representatives of the genus Pholadidea are found in the temperate and tropical por-
tions of all seas other than the Western Atlantic.

Genotype, Pholadidea loscombiana Turton, monotypic.

Key to the subgenera of Pholadidea

1. Umbonal reflections raised well above the umbos, mesoplax
small and lacking a basal portion, siphonoplax cup-like,
siphonal tube lacking. . . . .. . . . . Pholadideas.s.

2. Umbonal reflection closely appressed over the umbos, meso-
plax with a basal portion, siphonoplax variable, siphonal tube
PRCSOME Ray ts Sue are Se Deol eB la ae at eee

Subgenus Pholadidea Turton

Pholadidea ‘Goodall’ Turton 1819, A Conchological Dictionary of the British Islands, London, p. 147.
Talonella Gray 1851, Annals and Magazine Natural History (2) 8, p. 385 (genotype, Talona tridens Gray,
monotypic).

Shell small to medium in size, oval in outline, divided into two regions by an umbonal-
ventral sulcus, strongly beaked and gaping anteriorly in the young stage and producing
a callum when adult. Valves rounded posteriorly when young, truncate in the adult stage
and terminating with a cup-like siphonoplax. Siphonoplax entirely chitinous or with in-
ternal calcareous plates. Siphonal tube lacking. Umbonal reflections free for their entire
length. Protoplax lacking. Mesoplax very small and composed of two more or less tri-
angular pieces. Metaplax and hypoplax lacking.

Species in this subgenus, so far as known, are found only in the Eastern Atlantic and
the Indo-Pacific. However, a description of the subgenus and of Pholadidea loscombiana
Turton are included for a complete understanding of the genus.

Subgenotype, Pholadidea loscombiana Turton, monotypic.

90 JOHNSONIA, No. 34 Pholadidea

Pholadidea (Pholadidea) loscombiana Turton
Pholadidea loscombiana “Goodall,’ Turton 1819, A Conchological Dictionary of the British Islands, p. 147

(Exmouth [England] ).

Pholas papyraceus Solander 1786, A Catalogue of the Portland Museum, p. 82, lot 1828 [nomen nudum |.

Pholas papyraceus Turton 1822, Conchylia Insularum Britannicarum, p. 2, pl. 1, figs. 1-4 (Torbay [Eng-
land | ).

Pholas lamellata Turton 1822, Conchylia Insularum Britannicarum, p. 4, pl. 1, figs. 5-6 (Torbay [England] ).

Pholas striata de Blainville 1825, Manuel de Malacologie, p. 578, pl. 80 [bis] fig. 7; non Pholas striata
Linné 1758.

Pholas goodall {sic| de Blainville 1826, Dictionnaire des Sciences Naturelles 39, p. 532.

Pholas lamellosus ‘Turton’ Sowerby 1834, Proceedings Zoological Society London, p. 69 [error for lamellata
Turton].

Pholas vibonensis Philippi 1844, Enumeratio Molluscorum Siciliae 2, p. 4, pl. 13, fig. 5 (near Monteleone,
Sicily).

Cadmusia solanderia Leach 1852, A Synopsis of the Mollusca of Great Britain, p. 254, pl. 11, figs. 1-2.

IOS

Pholadidea papyracea var. aborta Jeffreys 1865, British Conchology 3, p. 117.
Pholadidea loscombeana ‘Goodall’ Lamy 1926, Journal de Conchyliologie 69, p. 138 [error for loscombiana

Turton].

Pholadidea solanderiana ‘Leach’ Lamy 1926, Journal de Conchyliologie 69, p. 140 [error for solanderia Leach].

Distinctive characters. Shell reaching about 14 inches in length. Umbonal reflections .
narrow and raised well above the umbos, mesoplax very small and in two parts. Shell
truncate posteriorly in the adult stage. Siphonoplax thin, chitinous and with the two
parts partially fused to form a tube. There is no siphonal tube.

Description. Shell reaching 31 mm. (about 14 inches) in length and 17 mm. (about ?
inch) in height, light in structure and producing a callum in the adult stage. Immature
specimens strongly beaked anteriorly and widely gaping, tapering to a rounded, posterior
margin. Valves divided into two regions by a narrow umbonal-ventral sulcus. Anterior
portion sculptured with concentric ridges and rather weak radial ribs. Rounded imbrica-
tions are produced where the concentric ridges and radial ribs cross one another. Poste-
rior portion sculptured by rather strong concentric ridges and growth lines. Umbonal

Plate 55. Pholadidea loscombiana Turton. English Channel, England. All adult specimens (1.4x). Fig. 1.
Dorsal view showing the small mesoplax and the cup-like siphonoplax. Fig. 2. Side view showing the
siphonoplax and the truncation of the valve posteriorly in the adult stage. Fig. 3. Inner view of valve
showing the small apophysis, the rather large chondrophore and muscle sear. Fig. 4. Side view showing
the extent to which the callum may extend anterior to the beaks.

Eastern Atlantic JOHNSONIA, No. 34 91

reflections in young specimens simple, narrow and raised well above the surface of the
shell. Pedal gape in adult specimens closed by athin callum which protrudes beyond the
beaks and is sculptured by faint growth lines. When the callum is complete only a nar-
row slit remains between the two halves and this is covered with a heavy periostracum,
only a minute pore remaining open. The callum is continued dorsally over the umbonal
reflection where it recurves upon itself and encloses the anterior adductor muscle. Proto-
plax lacking. Mesoplax small and composed of two triangular pieces. Metaplax and hy-
poplax lacking. Rounded posterior margin of the young specimen becoming truncate in
the adult stage, forming the attachment area for the thin, chitinous, cup-like siphonoplax.
Periostracum and siphonoplax light straw-yellow in color.

Interior of the shell white and usually glazed. Umbonal-ventral sulcus evident intern-
ally as a pronounced ridge. Muscle scars well marked. Pallial sinus broad and deep, ex-
tending anteriorly to the umbonal-ventral ridge, but often barely visible. Apophyses
small, fragile and projecting anteriorly from beneath the umbos at a sharp angle. A
rather large chondrophore is present on the left valve, a smaller one on the right.

Siphons united and capable of extending two to three times the length of the shell.
They are a pale to rather dark reddish-brown in color with a narrow band of white at the
posterior extremity. Incurrent siphon nearly twice the diameter of the excurrent siphon
and fringed with a few large and several small cirri. Exhalant siphon lacking cirri. Just
anterior to the siphonal opening there is a ring of white cirri surrounding both siphons.
Foot in young specimens more or less oval in outline and truncate, becoming atrophied
in the adult. Foot and mantle white. Forbes and Hanley (1853, 1, pl. F, fig. 4) figure
in color the siphons of a living specimen.

length * height ratio h:]
34.5 mm. 15.0 mm. 2.3 English Channel, England
30.0 16.0 1.8 * rs i
26.0 1550 Lot Plymouth, England
23.5 15.0 1.5 on -
21.0 11.0 1.9 - ze

* All specimens measured had a complete callum

Types. The holotype of P. loscombiana Turton is probably in the British Museum
(Natural History). The single specimen on which the name was based was in the Los-
combe collection, some of which was later obtained by Sowerby. The location of the
types of P. lamellata Turton is unknown. According to J. D. Dean (1936, Journ. of
Conchology 20, p. 233) the Turton collection was acquired by J. G. Jeffreys and with
the Jeffreys collection later went to the United States National Museum. Dr. Rehder,
however, informs me that they are not in Washington though a cotype of P. papyraceus
Turton is there.

Remarks. Pholadidea loscombiana Turton is the only species in European waters which
produces a callum in the adult stage and so is easily recognized. It appears to be most
closely related to Pholadidea tridens Gray, a New Zealand species, but differs from this
latter species by having a thin and entirely chitinous siphonoplax without any internal
calcareous plates. The young of P. loscombiana have been confused with Zifaea crispata
Linné, but differ by having a pronounced umbonal-ventral sulcus and by being closed
posteriorly. In addition, the siphons of the two species are very different.

92 JOHNSONIA, No. 34 Pholadidea

Pholadidea loscombiana Turton is restricted in its range to the British Isles and the
coast of France probably as far south as northern Spain, but records outside of the British
Isles are rare. They bore into mud, clay, peat, waterlogged wood and sandstone and are
usually found at low tide level or below, such specimens having been dredged by fisher-
men off the coast of Ireland. The wide variety of substrata in which this species lives is
no doubt responsible for much of the variability of the species and the large number of
synonyms noted above.

Range. Ireland, southern England, and south along the coast of France to San Sebas-
tian [Spain] (Lamy, 1926, p. 141).

Specimens examined. ENGLAND: Torquay (Redpath Museum, McGill University);
Plymouth (F. S. Russell); English Channel (MCZ). France: Cape Breton; Bay of
Biscay (both USNM).

Plate 56. Pholadidea loscombiana Turton. English Channel, England. Fig. 1. Young specimen showing pedal
gape and hinge area, the apophysis on the left valve is lost but the chondrophore is evident. Apophysis
showing on the right valve. Fig. 2. Side view of young specimen to show the high umbonal reflection and
the rounded posterior margin (both about 3.3x).

Subgenus Hatasia Gray
Hatasia Gray 1851, Annals and Magazine of Natural History (2) 8, p. 385.

Shell small to medium in size, oval to elongate-oval in outline, divided into two regions
by an umbonal-ventral sulcus, and producing a callum in the adult stage. Shells strongly
beaked and widely gaping anteriorly in the young stage, rounded and closed posteriorly ;
the siphon being capable of complete retraction within the shell. Siphonoplax chitinous.
Siphonal tube calcareous. Protoplax lacking. Mesoplax in the young stage small, flat,
more or less semicircular in outline and in one piece. In the adult stage an upper surface
is added to the mesoplax covering the posterior portion of the anterior adductor muscle.
This upper surface is divided into two parts longitudinally and is usually inset on the
young basal portion. Incipient metaplax and hypoplax present or absent.

Subgenotype, Pholas melanura Sowerby, subsequent designation, Stoliczka 1870.

Key to the species of Hatasia

1. Siphonoplax tubular with swellings at the base ..... 2
Siphonoplax diverging, without swellings at the base . tubifera

©

Posterior portion of valves inflated, shell generally over 28 mm.
in length, periostracum and siphonoplax nearly black . melanura
Posterior portion not inflated, shell generally under 20 mm. in
length, periostracum and siphonoplax light brown . . quadra

Eastern Pacific JOHNSONIA, No. 34 93

Pholadidea (Hatasia) melanura Sowerby
Plate 57

Pholas melanura Sowerby 1834, Proceedings Zoological Society London, p. 70 (Montem Christi, Columbiae
Occidentalis [Montecristi, Ecuador] ).

Penitella wilsonii Conrad 1849, Proceedings Academy Natural Sciences Philadelphia 4, p. 156 (Lower Cali-
fornia); Conrad 1850, Journal Academy Natural Sciences Philadelphia (2) 1, p. 279, pl. 39, fig. 4 (not fig. 5
as given in the text]; Conrad 1854, Journal Academy Natural Sciences Philadelphia (2) 2, p. 335.

Pholadidea (Hatasia) melanura Sowerby, Gray 1851, Annals and Magazine Natural History (2) 8, p. 385.

Pholadidea melanura Sowerby, Carpenter 1855, Reigen Collection of Mazatlan Mollusca, p. 8: P. Fischer
1858, Journal de Conchyliologie 7, p. 51; Fischer 1860, ibid. 8, p. 5, pl. 3, figs. 1-8.

Distinctive characters. Shell reaching about 24 inches in length. Posterior slope in-
flated. Basal portion of the siphonoplax with subreniform swellings, siphonoplax tubular.
Siphonal tube generally not firmly attached to the siphonoplax. Periostracum and si-
phonoplax nearly black.

Description. Shell reaching 56 mm. (24 inches) in length and 28 mm. (about 1 inch)
in height, thin, inflated, oval in outline and producing a callum in the adult stage. Im-
mature specimens beaked and widely gaping anteriorly, rounded and closed posteriorly.
Shell divided into two distinct regions by a pronounced umbonal-ventral sulcus. Anterior
portion sculptured with close-set, upturned, undulating ridges and radial ribs. Posterior
portion inflated and sculptured with rather strong concentric ridges. Umbos prominent
and located near the center of the dorsal margin. Umbonal reflections thin, simple, and
closely applied or raised just slightly above the surface of the umbos. Pedal gape in the
adult closed by a thin callum which protrudes anteriorly beyond the beaks and extends
dorsally over the umbonal reflections to enclose the anterior portion of the anterior ad-
ductor muscle. The callum is marked with faint growth lines and weak parallel ridges.
Mesoplax in young specimens thin, nearly flat, largely chitinous and semicircular in out-
line. The dorsal portion of the mesoplax is much narrower than the young basal portion

Plate 57. Pholadidea melanura Sowerby. Fig. 1. Panama Bay, 1 mile off the canal entrance in 10 fathoms.
Internal view of shell showing the small apophysis. Figs. 2-3. Penitella wilsoni Conr. (= Pholadidea melanura
Sowerby) Baja California. Fig. 2. Paratype, dorsal view showing the mesoplax with its large basal portion.
Fig. 3. Lectotype, lateral view showing the high umbonal reflection and the swelling at the base of the
siphonoplax (all natural size).

O4 JOHNSONIA, No. 34 Pholadidea

and is divided into two parts longitudinally. An incipient metaplax is formed by the
impregnation with calcium of the periostracum between the valves. Siphonoplax con-
sisting of a dark brown to nearly black chitinous tube. Adjacent to the valves there are
two subreniform swellings which fuse together to form the extended tubular siphonoplax.
Periostracum dark brown to black. Interior of shell white and usually glazed. Umbonal-
ventral sulcus evident internally as a strong, beaded ridge. Muscle scars well marked,
pallial sinus broad and deep, extending anteriorly beyond the umbonal-ventral ridge and,
in some specimens, nearly to the anterior margin of the shell. Apophyses small, solid,
slightly grooved at the free end and extending anteriorly from beneath the umbos at a
sharp angle.

We have not seen the soft parts of this species, but Fischer (1860, op. cit.) has figured
a specimen which he received from Panama.

length* height ratio h:]
56.0 mm. 28.0 20 Lectotype, P. wilsonii Conrad
aya 28.5 1.9 Paratype, . i
52.0 28.0 1.8 Panama Bay
48.0 26.0 1.8 fe
30.0 15.0 2.0 Paratype, P. wilsonii Conrad
28.0 6.5 4.0 ¥ a i

* All specimens measured had a complete callum

Types. The holotype of Pholas melanura Sowerby is inthe British Museum according
to a letter from G. L. Wilkins. The type locality is Monte Cristi, West Columbia
[Montecristi, Ecuador]. The lectotype of Penitella wilson Conrad is in the Academy of
Natural Sciences Philadelphia, no. 51012, from Baja California, Dr. Thomas B. Wilson
collector.

Remarks. Pholadidea melanura Sowerby is a very distinctive species, readily differen-
tiated from all others in this group by its nearly black periostracum and siphonoplax, and
by its inflated posterior portion. From Pholadidea quadra Sowerby, the species with
which it is most closely related, it differs by being much larger, having a much heavier,
darker periostracum and a larger siphonoplax.

This species is apparently rare and restricted in its distribution. We have seen only
five lots and two were from the same locality. The specimens collected by W. D.
Clarke off Panama were in 10 fathoms and it may well be that this species occurs abun-
dantly at such depths. However, as they are deep borers they seldom are brought up in
dredges and the shells remain in situ when the animal dies so that beach specimens are
practically unknown.

Range. From Baja California south to Ecuador.

Specimens evamined. Mexico: Baja California ( ANSP). Panama: Puerto Armuelles;
Guanico (both A. A. Olsson); Panama Bay, 1 mile off the Canal entrance in 10 fathoms
(W. D. Clarke). Ecuapor: San Pedro, near Manglaralto (A. A. Olsson).

Eastern Pacific JOHNSONIA, No. 34 95

Pholadidea (Hatasia) quadra Sowerby
Plate 58
Pholas quadra Sowerby 1834, Proceedings Zoological Society London, p. 71 (Montem Christi, Columbiae
Occidentalis [Montecristi, Ecuador] ); Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, p. 499, pl. 106,
figs. 62-63.
Pholadidea ( Hatasia) quadra Sowerby, Gray 1851, Annals Magazine Natural History (2) 8, p. 385.

Distinctive characters. Shell generally under 1 inch in length, oval in outline. Poste-
rior slope not inflated. Basal portion of siphonoplax with small oval swellings, siphono-
plax tubular, siphonal tube not firmly attached to the siphonoplax. Periostracum light
brown in color.

Description. Shell white, thin, fragile, broadly oval to quadrate in outline, reaching
19 mm. (# inch) in length and 10 mm. (about $ inch) in height and producing a callum
in the adult stage. Shell strongly beaked and widely gaping anteriorly in the young stage,
broadly rounded and closed posteriorly. Valves divided into two regions by a narrow and
only slightly impressed umbonal-ventral sulcus. Anterior portion sculptured with con-
centric ridges and radial ribs, the ribs being expressed mainly by the radial arrangement
of the rows of imbrications. Posterior slope sculptured with weak concentric ridges and
faint growth lines. Umbonal reflections thin, simple, rather wide and very closely ap-
pressed so that the sculpture below may show through. Pedal gape in adult specimens
closed by a large, thin callum which protrudes anteriorly well beyond the beaks. It is
sculptured with fine concentric growth lines and weak parallel ridges which are extensions
of the radial ribs. The callum extends dorsally over the umbonal reflection and encloses
the anterior adductor muscle. Mesoplax unknown except as figured by Sowerby. It is
more or less rectangular in outline with a wide basal portion and the upper part divided

Plate 58. Pholadidea quadra Sowerby. Figs. 1-2. Ecuador (about 3x). Fig. 1. Interior view of adult showing
the minute apophysis, the pronounced muscle sears and the pallial sinus. Fig. 2. External view showing
the greatly produced callum, the siphonoplax and the beginning of the siphonal tube. Figs. 3-4. Copies
of the figure from Sowerby, Thesaurus Conchyliorum 2, pl. 106, figs. 62-63 (2.2x). Fig. 8. Dorsal view
showing the divided mesoplax with its large basal area, the dorsal extension of the callum and the siphon-
oplax. Fig. 4. Side view showing the siphonal tube. The dorsal extension of the callum is lacking in both
figures 2 and 4.

96 JOHNSONIA, No. 34 Pholadidea

longitudinally into two parts. Metaplax and hypoplax lacking. Siphonoplax chitinous,
light brown in color, and with swellings near the base. Siphonal tube calcareous, thin
and firmly attached. Periostracum thin, a light red-brown in color and persistent. In-
terior of shell white and glazed. Umbonal-ventral sulcus evident internally as a beaded
rib. Muscle scars well marked. Pallial sinus broad and deep, extending anteriorly to
beyond the umbonal-ventral rib. Both concentric and radial sculpture visible internally.
Apophyses very small, solid, and extending from beneath the umbo anteriorly at a
rather sharp angle.
The soft parts of this species are unknown.
length height ratio h:]
19mm. 10mm. Lo Ecuador

Types. The location of the types of Pholas quadra is unknown. They are not in the
British Museum according toa letter received from G. L. Wilkins. A possible cotype is
in the P. P. Carpenter collection in the Redpath Museum. The type locality is Monte
Christi, West Columbia { Montecristi, Ecuador].

Remarks. We know practically nothing concerning Pholadidea quadra Sowerby, ex-
cept that it does exist. It was originally collected by Hugh Cuming in loose sand and
clay at Montecristi, Ecuador. It is most closely related to P. melanura Sowerby from
which it differs by being much smaller, having a less inflated posterior portion, and a
light brown rather than nearly black periostracum. From P. tubifera it differs by having
a tubular, rather than diverging, siphonoplax, and in having swellings at the base of the
siphonoplax.

Range. Known only from Ecuador.

Specimens examined. Ecuapor: (MCZ: Redpath Museum): Crucita, east of Manta
(A. A. Olsson).

Pholadidea (Hatasia) tubifera Sowerby
Plate 59

Pholas tubjfera Sowerby 1834, Proceedings Zoological Society London, p. 71 (Sinum Caraccensem, Colum-
biae Occidentalis [Bahia de Caraques, Ecuador] ).

Pholas tubifer Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, p. 499, pl. 106, figs. 64-65; Sowerby
1849, Proceedings Zoological Society London, p. 162, pl. 5, fig. 5 [error for tubjfera Sowerby |.

Pholadidea ( Hatasia) tubjfera Sowerby, Gray 1851, Annals Magazine Natural History (2) 8, p. 385.

Pholadidea tubjfera Sowerby, H.and A. Adams 1865, Genera of Recent Mollusca 2, p. 329; Fischer 1853,
Journal de Conchyliologie 7, p. 243; Dall 1909, Proc. United States National Museum 37, no. 1704, p. 277.

Distinctive characters. Shell reaching about 14 inches in length and elongate-oval in
outline. Basal portion of the siphonoplax diverging, siphonal tube heavy, calcareous and
firmly attached to the siphonoplax.

Description. Shell elongate-oval in outline, reaching 37.5 mm. (about 14 inches) in
length and 14 mm. (about $ inch) in height, solid in structure and producing a callum
in the adult stage. Immature specimens beaked anteriorly and widely gaping: narrowly
rounded posteriorly and closed. Shell divided into two regions by a narrow and often

Eastern Pacific JOHNSONIA, No. 34 97

weak umbonal-ventral sulcus. Anterior portion sculptured with very close-set, laminated
undulating ridges and radial ribs. The radial ribs are expressed only by the radial arrange-
ment of the undulations and the thickening of the ridges where the ribs cross. Umbonal
reflections simple, narrow and usually appressed. Pedal gape in adult specimens closed
by arather heavy, protruding callum which extends anteriorly well beyond the beaks.
The callum continues dorsally over the umbonal reflection and forms the enclosure for
the anterior portion of the anterior adductor muscle. Mesoplax in young specimens un-
known. In adult specimens it is more or less square to rectangular in outline and divided
into two parts longitudinally. Metaplax and hypoplax lacking. Siphonoplax composed
of diverging, chitinous flaps with a heavy fold posteriorly which unites the two parts and
forms the base to which the long calcareous siphonal tube is attached. Periostracum thin,
light brown in color and persistent.

Interior of shell white and usually glazed. Umbonal-ventral sulcus evident internally
as arib. Muscle scars well marked. Pallial sinus broad and deep, extending anteriorly to
just beyond the umbonal-ventral ridge. Concentric sculpture clearly visible internally.
Apophyses very small and weak, and extending from beneath the umbo anteriorly at a
rather sharp angle.

We have not seen the soft parts of this species, but Fischer (1858, Journal de Con-
chyliologie 7, p. 247) gives a description of specimens which he received from Panama.

length height ratio h:]
387.5 mm. 14.0 mm. 2.6 Panama City, Panama
22.0 EO 2.0 Panama
22.0 10.0 2.2 ¥
20.0 8.0 2.5 -
15.0 825 its i

Types. The types of Pholas tubifera Sowerby are in the British Museum (Natural
History) according to a letter received from G. L. Wilkins. The type locality is Bahia
de Cardques, Ecuador.

Remarks. Pholadidea tubifera Sowerby is not closely related to any species of Phola-
didea in the Eastern Pacific. It approaches P. guadra in size, but is generally a much

Plate 59. Pholadidea tubifera Sowerby. Panama. Fig. 1. Dorsal view showing the divided mesoplax, diverging
siphonoplax and siphonal tube. Fig. 2. Side view showing protruded callum. Fig. 3. Ventral view. Fig.
4. Internal view of valve to show the umbonal-ventral ridge, muscle scars and pallial sinus (all about 2x),

98 JOHNSONIA, No. 34 Lignopholas

heavier shell, has a diverging rather than tubular siphonoplax and a heavy siphonal tube.
See also remarks under P. guadra Sowerby.

Little is known concerning this species. Sowerby states in the original description that
Hugh Cuming obtained the specimens from a piece of decayed wood dredged in 10 fath-
oms. Dr. Olsson writes that the specimen collected at Esmeraldas, Ecuador were in shale.
The Hassler obtained several fine specimens from off Panama and it may well be that in
certain restricted areas they are abundant.

Range. Panama south to Payta, Peru (Dall 1909).

Specimens examined. PaNaMa: Balboa, Canal Zone; Panama City (both J. Zetek);
off Panama (Hassler Voyage). Ecuapor: Esmeraldas (A. A. Olsson).

Genus Lignopholas, new genus’

Species in this genus are small wood borers. They are elongate-oval to pear-shape in
outline, the valves are divided into two distinct areas by an umbonal-ventral sulcus and
a callum is produced in the adult stage. Shells beaked and widely gaping anteriorly in
the young stage; the beaks being truncated give the shell a teredo-like appearance. Valves
narrowly rounded and closed posteriorly. Posterior slope covered by a series of overlap-
ping, thin, chitinous lamellae which are fringed on their posterior margin. Umbos promi-
nent and located near the center of the dorsal margin in young shells, but in the adults
they are very near the anterior end due to the disproportionate growth of the posterior
slope. Mesoplax divided longitudinally; all other accessory plates Jacking. Sulcus ex-
pressed internally as a pronounced umbonal-ventral ridge. Apophyses very long, thin,
and fragile. There is a pronounced chondrophore in the left valve and an internal ligament.

The species in this genus superficially resemble Martesia, particularly in the young
stage. They have the funnel-shaped pit below the umbonal reflection and a similarly shaped
pedal gape. However, the adults are readily distinguished by the lack of both the meta-
plax and the hypoplax, and in addition, by the presence of the chitinous lamellae on the
posterior slope.

There are only two known species in this genus. One, L. rivicola Sowerby is a fresh-
water species recorded only from Borneo; the other, L. clappi Turner, is known only
from Bluefields, Nicaragua, where it occurs in brackish water. Both species appear to
bore into wood near the surface of the water rather than at the mud line as do most pho-
lads. It is impossible at this time to explain the distribution of these two closely related
species.

Little is known concerning the destructiveness of L. rivicola Sowerby, but it is re-
corded as boring into floating timbers of the landing pier at Gunung Tebur, Borneo. At
Bluefields, Nicaragua, L. clappi appeared to be rather destructive during the period that
test boards were submerged there.

'From the Latin lignum (wood) and pholas.

Western Atlantic JOHNSONIA, No. 34 99

Lignopholas clappi, new species !
Plate 60
Distinctive characters. A small wood borer superficially resembling a Martesia, but
lacking the metaplax and hypoplax, and having the mesoplax divided longitudinally into
two parts. In addition, there is a series of overlapping chitinous lamellae on the poste-
rior slope.

Description. Shells variable in size and shape, the larger specimens reaching about 25
mm. (1 inch) in length and about 6 mm. (+ inch) in height. Specimens generally a slen-
der pear-shape in outline, white to light brown in color and having a callum in the adult
stage. Young specimens are beaked and widely gaping anteriorly; the beaks being trun-
cated, give the shell a teredo-like appearance. Posteriorly the shell tapers to a rounded
margin which may be broad or narrow depending largely upon the age of the specimen.
Valves divided into two distinct regions by an umbonal-ventral sulcus. Anterior portion
sculptured by close-set denticulated ridges. Posterior portion sculptured with smooth,
rounded, concentric ridges which are continuations of the denticulated ridges of the an-
terior slope. In addition, the posterior portion has a series of overlapping chitinous lam-
ellae which are coarsely fringed. An adult specimen may have six to eight of these lamel-
lae. Umbos prominent, located near the center of the dorsal margin in young specimens,
but in adults, very close to the anterior end due to the disproportionate growth of the

a TRS oon
Se

Plate 60. Lignopholas clappi Turner. Fig. 1. Side view of the holotype showing the extent to which the meso-
plax extends over the beaks and the chitinous lamellae on the posterior slope. Fig. 2. Dorsal view of a para-
type to show the divided mesoplax and the lack of the metaplax. Fig. 3. Internal view of a young speci-
men. Fig. 4. External view of a young specimen. Fig. 5. Dorsal view of the mesoplax of a young specimen.
Fig. 6. Dorsal view of the mesoplax of an adult specimen. Fig. 7. Ventral view. Fig. 8. Side view. All
from Bluefields, Nicaragua. Figs. 1-2 (4x); Figs. 3-8 (6x).

1Named for the late Dr. William F. Clapp, of the Clapp Laboratories, Duxbury, Massachusetts.

100 JOHNSONIA, No. 34 Lignopholas

posterior portion. Umbonal reflections rather small, thin and closely appressed over the
umbos. A rather heavy callus extends over the umbo where the anterior adductor mus-
cle is attached. In adult specimens the pedal gape is closed by a thin, smooth callum
which does not extend dorsally between or over the beaks. The two halves of the com-
pleted callum are connected by a thin periostracum, only a minute pore remaining open
in the fully adult specimen. Protoplax lacking. Mesoplax in young specimens more or
less semicircular in outline with a median groove and faint concentric growth lines. In
the adult, the dorsal portion of the mesoplax grows forward, completely covering the an-
terior adductor muscle and extending over the beaks. Interior of the shell white and
glazed. Muscle scars only faintly indicated, the large oval posterior adductor muscle scar
set high on the posterior slope. Pallial sinus extending anteriorly to the umbonal-ventral
ridge. Periostracum thin, light straw-yellow and persistent. The soft parts of this spe-

cies are unknown.

length* height ratio h:]
24.2 mm. 7.0mm. 3.4 Holotype
18.0 6.3 2.8 Paratype
12.8 5.4 2.3 ¥.
15.0 3.8 3.9 re
5.0 2.8 Ne se

* All specimens measured had a complete callum

Types. Holotype, Museum of Comparative Zodlogy, no. 200,046 from Bluefields,
Nicaragua. Paratypes from the same locality are in the collection of the Museum of
Comparative Zoélogy and the United States National Museum.

Remarks. This is a very distinctive species most closely related to L. rivicola Sowerby
from freshwater at Gunung Taboor [Gunung Tebur], twelve miles up the Pantai River
in Borneo, a species collected during the voyage of the Samarang. Lignopholas clappi is
a smaller, more slender species with a much finer fringe on the chitinous lamellae and in
addition, the mesoplax of clappi is pointed posteriorly rather than truncate. It is a wood
borer and is found in both brackish and marine conditions. Of our American species, ZL.
clappi superficially appears close to species of Martesia, but it lacks both the metaplax
and the hypoplax, and in addition, has chitinous lamellae on the posterior slope, a con-
dition not found in Martesia.

Specimens of this species were taken from test boards suspended from the wharf in
Bluefields Lagoon at a depth of eighteen inches below the surface. At this station the
water was brackish and sluggish. he board was submerged under the supervision of the
W. F. Clapp Laboratories, Duxbury, Massachusetts, on June 12, 1985 and removed
February 28, 1936. A second test board was submerged at Schooner Cay, Bluefields,
Nicaragua, on December 27, 1985 and removed April 10, 1986. At this locality the
water though still a little brackish was far more affected by the tides. Young and adult
specimens were taken from both boards which would indicate that breeding probably
occurs throughout the year.

It is curious, that despite all the collecting and test board studies which have been made
along the Central American coast, this species has been taken only in this one area.

Western Atlantic JOHNSONIA, No. 34 101

It is apparently abundant at Bluefields, judging from the numbers that occurred in the
test boards.

Range. Known only from the type locality.

Specimens examined. NicAraGua: Schooner Cay and Bluefields Lagoon, Bluefields

(MCZ; USNM).

Genus Martesia Sowerby

Martesia ‘Leach’ Sowerby 1824, Genera of Recent and Fossil Shells, part 23, Pholas, p. 2 (genotype, Pho-
las clavata Lamarck {= M. striata Linné], monotypic); de Blainville 1825, Manuel de Malacologie, p. 632.
Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104, no. 11, p. 2.

Mactresia Gray 1840, Synopsis Contents British Museum, ed. 42, p. 154 [error for Martesia Sowerby }.

Mactesia ‘Gray’ Paetel 1875, Familien-und Gattungsnamen Mollusken, p. 119 [error for Martesia Sowerby }.

Shells rather small, not reaching over 50 mm. (2 inches) in length, light in structure,
white to light ivory in color, generally pear-shaped in outline, divided into two distinct
regions by an umbonal-ventral sulcus and producing a callum in the adult stage. Shells
beaked and widely gaping anteriorly in the young stage; the beaks sinuously to sharply
truncated, giving the shells a teredo-like appearance. Valves narrowly to broadly rounded
posteriorly and closed. Siphons capable of complete retraction within the shell. Umbos
prominent, nearly centrally located in young shells. In the adult they are very near
the anterior end due to the disproportionate growth of the shell. Umbonal reflections
closely appressed over the umbos but free anteriorly. They are broadly recurved, form-
ing a funnel-shaped pit below. Protoplax lacking. Mesoplax variable, ranging from cir-
cular to cuneiform in outline. Metaplax and hypoplax long, narrow and pointed anteri-
orly. Posteriorly they may be pointed, truncate or divided. Interior of the shell with a
pronounced umbonal-ventral ridge which is enlarged to form a condyle at the ventral
margin. Chondrophore and internal ligament present, but small. Apophyses long, thin
and fragile.

The genus Martesia, like nearly all genera of the Pholadidae, is clear-cut and easily
distinguished. It is characterized by the teredo-like shell in the young stage, by the
presence of a mesoplax, metaplax and hypoplax, and by the funnel-shaped pit below the
umbonal reflection. In addition, the species normally bore only into wood, seeds or other
woody substances. The genus is probably most closely related to Diplothyra, which has
often been considered a subgenus of Martesia. See also under Diplothyra.

Species of Martesia occur throughout the world in nearly all temperate and tropical
regions, but their distribution is rather difficult to understand at present. Undoubtedly
a thorough knowledge of the life histories of the various species concerned will answer
many of the problems. Martesia striata Linné, the best known and most widely distrib-
uted species in the genus, is the most destructive and has succeeded in invading nearly
all tropical and south temperate seas. Whether this ability to spread is due to a wide
range of temperature and salinity tolerance or to some factor in the life history of this
species, it is impossible to say. However, Martesia cuneiformis Say, which is often found
living with M. striata Linné, is restricted to the Western Atlantic. A third species, JZ.
fragilis Verrill and Bush, appears to be a pelagic species, with only occasional specimens
being taken from fixed structures.

102 JOHNSONIA, No. 34 Martesia

Two factors complicate the taxonomy of this genus. The first is the great variability
within the species. This is an expression of the rate of growth and amount of crowding
of the specimens as well as the hardness of the substratum in which they are living. Speci-
mens living in overcrowded conditions may reach maturity and produce a callum when
not more than 5 mm. in length. Such specimens are often referred to as stenomorphs, a
term more generally used in the Teredinidae. Other specimens with ample room, boring
into soft wood such as palmetto, may reach a length of 40 mm. or more before producing
acallum. These specimens usually have thin, perfectly formed shells with widely spaced
rows of imbrications and a smooth, nearly circular mesoplax. However, equally un-
crowded specimens boring into hard, knotted wood usually have closely set rows of im-
brications; the shell may be variously curved to fit around obstructions in the burrow,
and the mesoplax is often badly deformed. From the large amount of material available
for study it is obvious that the number of rows of imbrications cannot be used as a char-
acter to distinguish species. The second factor complicating the taxonomy is the great
difference between the young and adult specimens. This has resulted in the description
of several genera based upon the young stages of the various species. As shown in the
diagram (plate 64) the valve of a young specimen is nearly circular in outline with a cen-
trally located umbo. Gradually the posterior portion elongates so that just before the
callum is produced the posterior portion may be two and one-half to three times as long
as the anterior portion. Usually at the time the callum is produced an unsculptured por-
tion is also added to the posterior end of the valve making it as much as five to six times
the length of the anterior portion.

Genotype, Pholas clavata Lamarck (= P. striata Linné), monotypic.

Two subgenera in the genus Martesia may be separated as follows:

1. Mesoplax oval to circular, metaplax and hypoplax pointed or truncate
posteriorly, netidivided...-. 2°25 7225 2 ae See
2. Mesoplax wedge-shaped or cuneiform, metaplax and hypoplax divided
posteriorly i: 8 oc. 459. La og Wat se a ee

Subgenus Martesia Sowerby

Martesia ‘Leach’ Sowerby 1824, Genera of Recent and Fossil Shells, part 23, Pholas, p. 2 (genotype, Pho-
las clavata Lamarck { = M. striata Linné], monotypic); de Blainville 1825, Manuel de Malacologie, p. 632.

Martesiella Verrill and Bush 1898, Proceedings United States National Museum 20, no. 1139, p. 777 (sub-
genotype, Martesia (Martesiella) fragilis Verrill and Bush).

Hiata Zetek and McLean 1936, Nautilus 49, p. 110 (genotype, Hiata infelix Zetek and McLean [=M.
striata Linné], original designation).

Mesopholas Taki and Habe 1945, Venus 14, p. 109 (genotype, Mesopholas intusgranosa Taki and Habe
[= M. striata Linné], original designation).

Diploplax Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104, no. 11, p. 10 (subgeno-
type, Martesia (Diploplax) americana Bartsch and Rehder | =M. striata Linné], original designation).

The species in the subgenus Martesia are characterized by having the mesoplax broadly
oval to circular in outline and having the metaplax and hypoplax pointed or truncate
and not divided. In addition, there is a sickle-shaped shield over the umbo which forms
the attachment area for the anterior adductor muscle. The shield is attached anteriorly,
but free posteriorly, and raised well above the surface of the umbo. The transverse, basal
portion of the mesoplax fits beneath the free end of the shield. The beaks are truncated

Western Atlantic JOHNSONIA, No. 34 103

nearly at right angles to the anterior margin of the disc, giving the shell a teredo-like
appearance.
This subgenus is distributed throughout the temperate and tropical regions of the world.

Subgenotype, Pholas clavata Lamarck (= M. striata Linné), monotypic.

Key to the American species of Martesia ss.

1. Mesoplax of adult specimens more or less circular, inflated and sculptured
only by itresular wrinkles = 20. 4... «.. « « = » «+ « « Mi striala
2. Mesoplax oval, dorsal portion depressed, with a peripheral keel and definite
COMGEMENIEISCONPEEEG |. 4 2s ow a) we a ee, 2 ME repli

Martesia (Martesia) striata Linné
Plates 35: 61-64

Pholas striata Linné 1758, Systema Naturae, ed. 10, p. 669 (Western Europe); Dodge 1952, Bulletin Amer-
ican Museum Natural History 100, Art. 1, p. 26.

Pholas pusillus Linné 1758, Systema Naturae, ed. 10, p. 670 (America); Dodge 1852, Bulletin American
Museum Natural History 100, Art. 1, p. 27.

Pholas conoides Parsons 1765, Philosophical Transactions 55, p. 1, pl. 1 [abridged edition 12, 1763-1769,
p. 174, pl. 5, figs. 3-6].

Pholas nanus “Solander’ Pulteney 1799, Catalogue of the Birds, Shells, ete. of Dorsetshire [in] Hutchins
History of Dorset County, p. 27 (in sides of ships at Poole and Waymouth [sic], England).

Pholas faleata Wood 1815, General Conchology, London, p. 84, pl. 16, figs. 5-7 [young].

Pholas clavata Lamarck 1818, Histoire Naturelle des Animaux Sans Vertébres 5, p. 446 (Seas of Western
Europe and America).

Pholas tenuistriata de Blainville 1826, Dictionnaire de Sciences Naturelles 39, p. 531 (Central America,
based on Encyclopédie Méthodique 1792, Atlas 2, pl. 170, figs. 4-8 [not pl. 169, figs. 4-8 as given by de
Blainville] ).

Pholas decussata “Valenciennes’ Bory de Saint-Vincent 1827, Tableau Encyclopédique et Méthodique Atlas
pe 145, pl. A770, faes. 1-3;

Plate 61. Martesia striata Linné. Fig. 1. Dorsal view of an adult specimen showing the nearly circular, in-
flated mesoplax and the long, narrow metaplax. Fig. 2. Ventral view showing the callum and the hypo-
plax. Fig. 3. Side view showing the thin, almost entirely chitinous posterior portion of the valve which is
characteristic of rapidly growing specimens. All specimens are from Trinidad, British West Indies taken
from a test board submerged July 16, 1948 and removed August 16, 1949 (all about 13x).

104 JOHNSONIA, No. 34 Martesia

Pholas atomus ‘Valenciennes’ Bory de Saint-Vincent 1827, Tableau Encyclopédique et Méthodique, Atlas
1, p. 145, pl. 170, figs. 4-8.

Pholas ovum Wood 1828, Supplement to the Index Testaceologicus, p. 2, pl. 1, fig. 4 (West Indies).

Pholas hornbeckii d’Orbigny 1842 [in] Sagra, Histoire Naturelle de ]’Ile de Cuba, Atlas, pl. 25, figs. 23-
25: d’Orbigny 1853, ibid., text, 2, p. 217 (St. Thomas, Hornbeck collector).

Penitella xilophaga Valenciennes 1846, Voyage Autour du Monde sur la Frégate La Vénus, Atlas de Zoolo-
aie, Mollusques, pl. 24, fig. 2; non Pholas xylophaga Deshayes 1835.

Pholas teredinaeformis Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, p. 490, pl. 108, figs. 97-98 (in
cake of floating wax off Cuba).

Pholas corticaria ‘Gray’ Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, p. 495, pl. 108, figs. 94-96 (no
locality given); [Bay of Port Royal, Kingston, Jamaica, C. B. Adams 1850, Contributions to Conchology, no.
5, pp. 75-79].

Pholas terediniformis Sowerby 1849 [1850] Proceedings Zoological Society London, p. 161 [error for P.
teredinaeformis Sowerby }.

Pholas rosea C. B. Adams 1850, Contributions to Conchology, no. 5, pp. 75-76 (Bay of Port Royal, near
Kingston, Jamaica); Clench and Turner 1950, Occasional Papers On Mollusks 1, no. 15, p. 338.

Pholas beauiana Recluz 1853, Journal de Conchyliologie 4, p. 49, pl. 2, figs. 1-3 (Guadeloupe).

Martesia striata Linné, Tryon 1862, Proceedings Academy Natural Sciences Philadelphia, p. 92.

Pholas beaniana |sic] “Recluz’ Paetel 1890, Catalog Conchylien-Sammlung 3, p. 3 [error for beauiana Recluz
1853].

Martesia curta ‘Sowerby’ Dall 1909, Proceedings United States National Museum 37, p. 161; non Pholas
curla Sowerby 1834.

Penitella xylophaga ‘Valenciennes’ Lamy 1921, Bulletin Muséum d’Histoire Naturelle, Paris 27, p. 179
(error for P. xilophaga Valenciennes].

Penitella incisa; P. silicula; P. lignivora; P. stillata; P. larvata; P. pisum and P. phaseolina all ‘Valenciennes’
Lamy 1921, Bulletin Muséum d’Histoire Naturelle, Paris 27, pp. 180-181. [These manuscript names of
Valenciennes were introduced needlessly by Lamy as synonyms of Martesia striata Linné]}.

Pholas cupula Yokoyama 1924, Journal of the College of Science, Imperial University of Tokyo 45, Art. 1,
p. 87, pl. 2, fig. 15 (fossil, Numa, Awa, Japan) [young]; Habe 1952, Genera of Japanese Shells, Pelecy-
poda, No. 3, p. 244.

Martesia striata tokyoensis Yokoyama 1927, Journal Faculty of Science, Imperial University of Tokyo. Sec-
tion 2, 1, pt. 10, p. 428, pl. 48, figs. 2-3. (fossil, Tabata, Tokyo, Japan); Habe 1952, Genera of Japanese
Shells, Pelecypoda, No. 3, p. 244.

Martesia pulchella Yokoyama 1932, Journal Faculty of Science, Imperial University of Tokyo, Section 2,
3, pt. 6, p. 238, pl. 2, fig. 5. (Fossil, Uryu, Ishikari Prov., Japan).

Hiata infelix Zetek and McLean 1936, Nautilus 49, p. 110 (Balboa, Canal Zone, Panama) [young].

Martesia hawaiiensis Dall, Bartsch and Rehder 1938, Bulletin B. P. Bishop Museum, no. 153, p. 205, pl.
52, figs. 1-7 (Pearl Harbor, Oahu, Hawaiian Islands).

Martesia (Diploplax) americana Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104, no.
11, p. 13, pl. 2, figs. 1-2; pl. 3, figs. 3-4 (Fort Dade, Florida) [young].

Martesia ( Diploplax) funisicola Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104, no.
11, p. I+, pl. 3, figs. 1-2, 13-14 (Lake Worth, Florida in lead cable) [young].

Mesopholas intusgranosa Taki and Habe 1945, Venus 14, p. 110 (Wakayama, Honshu, Japan); Habe 1952,
Genera of Japanese Shells, Pelecypoda, No. 3, p. 244.

Mesopholas nucicola Taki and Habe 1945, Venus 14, p. 110 (Tanabe Bay, Wakayama, Honshu, Japan).

Martesia intercalata ‘Carpenter’ Hertlein and Strong 1950, Zoologica 35, no. 19, p. 250; non Martesia in-
tercalata Carpenter 1857.

Distinctive characters. Shell under 2 inches in length, pear-shaped, beaked anteriorly
and widely gaping in the young stage, producing a callum when adult, Umbonal reflec-
tion with a funnel-shaped pit below and a sickle-shaped flange over the umbo. Mesoplax
broadly oval to circular, inflated and sculptured only with irregular wrinkles. Metaplax
and hypoplax long, narrow and pointed at both ends.

Western Atlantic JOHNSONIA, No. 34 105

Description. Shell white, variable in size and shape, larger specimens reaching 44 mm.
(about 1} inches in length and 19 mm. (about ? inch) in height. Generally more or less
pear-shaped in outline and producing a callum in the adult stage. Young specimens
beaked anteriorly and widely gaping. Beak truncated at a sharp angle to the anterior
margin of the dise, giving the shell a teredo-like appearance. Posteriorly the shell is closed
and broadly to narrowly rounded depending upon the age and rate of growth of the speci-
men. In very rapidly growing specimens the posterior slope may be greatly extended,
this portion being thin and almost entirely chitinous and nearly devoid of sculpture.
Valves divided into two distinct regions by a shallow but distinct umbonal-ventral sulcus.
Anterior portion sculptured with close-set, denticulated, concentric ridges and usually

Plate 62. Martesia striata Linné. Fig. 1. Ventral view of a young specimen showing the large pedal gape and
the foot. Punta Gorda, Florida (3x). Fig. 2. Dorsal view of young specimen with the anterior adductor
muscle removed to show the flanges over the umbos and the semicircular mesoplax in place, fitting beneath
the flanges. Fort Dade, Florida (3x). Fig. 3. Internal view of the valve of a young specimen to show the
narrow apophysis, the umbonal-ventral ridge and the posterior adductor muscle scar. Fort Dade, Florida
(3x). Fig. 4. External view of the valve of a young specimen to show the flange which is raised well above
the umbo, the rather coarse denticulated ridges of the anterior slope and the low concentric ridges of the
posterior portion (3x). Fig. 5. Dorsal view of the holotype of Hiata infelix Zetek and McLean (= Martesia
striata Linné, young) showing the chitinous covering of the anterior adductor muscle during the young stage
(13x). Fig. 6. Side view of the paratype of Hiata infelix Zetek and McLean (14x). Fig. 7. Side view of an
adult specimen with an elongate, thin unsculptured posterior portion which is often produced when speci-
mens grow very rapidly. This should not be confused with the siphonoplax as the metaplax and hypoplax
extend the entire length (about 24x).

106 JOHNSONIA, No. 34 Martesia

weak radial ribs over the beaks. Posterior portion sculptured with smooth, concentric
ridges. Umbos prominent and located nearly at the anterior end of the shell in adult
specimens. In very young specimens the umbos are centrally located, but as the speci-
mens grow the posterior slope elongates. Umbonal reflections rather small, thin, free
anteriorly and broadly reflected, forming a funnel-shaped pit below. A sickle-shaped
flange extends over the umbo. It is attached anteriorly, but free and raised well above
the surface of the shell posteriorly. In adult specimens the pedal gape is closed by a thin
but strong callum which does not extend beyond the beaks and is usually nearly devoid
of sculpture. The narrow slit remaining between the two halves of the completed callum
is closed by a thin sheath of periostracum leaving open only a minute anterior pore in
the fully adult specimen. Protoplax lacking. Mesoplax in young specimens transverse,
more or less semicircular in outline, and with a shallow U-shaped to deep V-shaped em-
bayment anteriorly, depending upon the age of the specimen. The mesoplax in young
specimens is divided into two parts by a median groove and is marked with concentric
growth lines. In adult specimens the mesoplax grows dorsally and anteriorly enclosing
the anterior adductor muscle. The completed mesoplax is large, inflated, nearly circular
in outline, often with a slight notch posteriorly and asmall point anteriorly. The dorsal
surface is sculptured only with irregular wrinkles. Metaplax pointed at both ends, long,
narrow and bent downward anteriorly to fit under the mesoplax. Hypoplax pointed at
both ends. Siphonoplax lacking.

Interior of the shell white to light yellow in color and usually glazed. Umbonal-ventral
sulcus expressed internally as a beaded ridge which forms a condyle at its ventral margin.
Apophyses long and thin. Muscle scars well marked, the anterior adductor muscle being
attached to the sickle-shaped flanges over the umbos. Pallial sinus broad and deep, ex-
tending anteriorly beyond the umbonal-ventral ridge. Periostracum thin, light straw-
yellow in color and persistent.

Siphons white to light ivory in color, minutely papillose and capable of extension only
about the length of the shell. Incurrent siphon about three times the diameter of the
excurrent siphon, the aperture fringed with papillae which extend internally as ridges.
Excurrent siphon lacking papillae. Foot and mantle white to light ivory in color. Foot
in young specimens large, nearly circular in outline and truncate. It is atrophied in the
adult. he description of the soft parts is based upon preserved material.

length* height ratio h:]
44.0 mm. 19.0 mm. 2.3 Trinidad, B.W.1I.
40.0 13.0 3.1 Sneads Island, Manatee River, Florida
30.0 DNAs 1.4 Sanibel Island, Florida
25.0 9.0 oN, San Juan, Puerto Rico
17.0 10.0 17 oe te ys 5
5.5 2.5 2.2 aie mas a 2
3.5 Das 1.4 Ribault Bay, Mayport, Florida

* All specimens measured had a complete callum

Types. According to Hanley (1855, p. 25) and Dodge (1952, p. 26) Linné did not
have a specimen of Pholas striata in his collection. His only reference was to Gualtieri
1742, Index 'Testarum Conchyliorum, pl. 105, fig. F. We here select the figure of Gual-
tieri to represent the type. The locality given by Linné was Europe. We here restrict

Western Atlantic JOHNSONIA, No. 34 : 107

the type locality to Kingston, Jamaica, from which locality we have a good series.
Hanley and Dodge also state that Linné did not have a specimen of Pholas pusillus but
based his description upon the figure of P. Brown 1756, The Civil and Natural History
of Jamaica, p. 417, pl. 40, fig. 11. The type locality is Jamaica. The types of Pholas
clavata Lamarck and Penitella wilophaga Valenciennes are in the Paris Museum accord-
ing to Lamy. The types of Pholas teredinaeformis and Pholas hornbeckii d’Orbigny are
in the British Museum. Paratypes of Pholas rosea C. B. Adams are in the Museum of
Comparative Zoédlogy, no. 155634. The holotype of P. corticaria ‘Gray’ Sowerby is in
the British Museum. The type locality of these last two named forms is Bay of Port
Royal, Kingston, Jamaica (see C. B. Adams 1850, Contributions to Conchology, no. 5,
pp. 75-76, and Clench and Turner 1950, Occasional Papers On Mollusks 1, no. 15, p.
338). The holotype of Hiata infelia Zetek and McLean is in the Museum of Compara-
tive Zodlogy, no. 100088. The holotype of Martesia hawatiensis Dall, Bartsch and Rehder
is in the United States National Museum, no. 484213, as is the holotype of Martesia
(Diploplax) americana Bartsch and Rehder, no. 573550 and Martesia (Diploplax) funisi-
cola Bartsch and Rehder, no. 573551. Paratypes of Mesopholas nucicola Taki and Habe
are in the Museum of Comparative Zodlogy, no. 194818, from Tanabe Bay, Wakayama,
Honshu, Japan. Idiotypes of Mesopholas intusgranosa Taki and Habe from Shirahama,
Wakayama, Honshu, Japan are also in the Museum of Comparative Zoélogy, no. 194824.

Remarks. Martesia striata Linné is a very variable but distinctive species. Complete
adult specimens can be distinguished readily from all other species in this genus by the

Plate 63. Martesia striata Linné. Figs. 1-6. Holotype of Martesia (Diploplax) americana Bartsch and Rehder
(= Martesia striata Linné, young). Fig. 1. Internal view of the left valve showing the flange over the umbo,
the apophysis, umbonal-ventral ridge and the posterior adductor muscle scar. Fig. 2. External view of the
same valve showing the funnel-shaped pit below the umbonal reflection. Fig. 3. Internal view of the right
valve; the flange over the umbo and the apophysis are both lost. Fig. 4. External view of the right valve
Fig. 5. Ventral view of the mesoplax. Fig. 6. Dorsal view of the mesoplax (all about 5x).

108 JOHNSONIA, No. 34 Martesia

large, inflated, often irregularly shaped but generally nearly circular mesoplax which is
sculptured only by irregular wrinkles. In specimens boring into hard wood or in over-
crowded conditions the mesoplax may be malformed, often having a rough irregular sur-
face and scalloped edges. The extensive synonymy given above reflects the great varia-
bility. We have seen type material of many of the synonyms, while others were suffi-
ciently well described and figured as to leave no doubt as to their position. The speci-
mens named and described as Martesia curta Sowerby by Dall (1909, op. cit.) and Mar-
tesia intercalata Carpenter by Hertlein and Strong (1950, op. cit.) have been examined
and in both cases they proved to be M. striata Linné.

This species is most closely related to M. fragilis Verrill and Bush. The latter species,
however, is generally smaller and more delicate, and has a depressed, broadly oval mes-

GE
)

Plate 64. Martesia striata Linné. Fig. 1. Outline sketches of a series of specimens from young (on right) to
nearly adult (on left) to show the change in shape with age. Magnification is not constant, the young speci-
men was only 2 mm. in length while the one on the left was 2} em. All specimens were taken from a
single test board from San Juan, Puerto Rico. Fig. 2. A series of dorsal plates to show the range in shape,
size and condition. The central figure is of the mesoplax from a specimen which had met obstructions in
its burrow and so was deformed. Specimens from Daytona Beach, Florida and Guantanamo, Cuba. Fig. 3.
Dorsal and ventral view of a mesoplax from a young specimen from St. Petersburg, Florida. Fig. 4. Ven-
tral view of the mesoplax from a paratype specimen of Hiata infelix Zetek and McLean (= Martesia striata
Linné). Fig. 5. Mesoplax from avery young specimen from San Juan, Puerto Rico to show the pronounced
central groove. Fig. 6. Mesoplax from a young specimen from Guantanamo, Cuba, with a very weak cen-
tral groove. Fig. 7. Dorsal view of a typical, fully-developed mesoplax to show the irregular sculpture and
the cushion-like appearance. Fig. 8. Ventral view of the same specimen. Fig. 9. Side view. Figs. 7-9.
From San Juan, Puerto Rico.

Western Atlantic JOHNSONIA, No. 34 109

oplax which is always notched posteriorly and is clearly sculptured with concentric rings.
The valves of M. striata and M. fragilis are generally difficult to distinguish. Those of
M. fragilis, however, are usually shorter, more broadly rounded posterior to the umbos
and more uniformly sculptured. The valves of both species are distinguished from others
in the genus by the sickle-shaped flange over the umbo. The young of the two species
are impossible to distinguish unless the mesoplax is present. The dorsal portion of the
mesoplax in M. striata does not develop fully until the callum is being produced, whereas
the mesoplax of M. fragilis develops much earlier and may be complete in a half grown
specimen. The very young of the two species are completely indistinguishable, at least
in preserved material.

It is surprising that so little is known concerning the biology of this species consider-
ing its economic importance in certain localities. At San Juan, Puerto Rico, Cavite,
Luzon, Philippine Islands, and Pearl Harbor, Honolulu, Hawaiian Islands, this species
probably equals in its destructiveness the damage caused by the Teredinidae. From one
test board measuring 12’’x6’’x1"’ submerged at San Juan, Puerto Rico for eight months,
we obtained 208 specimens, many of which measured over 20 mm. in length. Test board
records at Guantanamo, Cubaand San Juan, Puerto Rico, indicate that this species prob-
ably breeds throughout the year. This is perhaps one explanation for its wide distribu-
tion in the tropics. It reaches maturity in one month and specimens four months old may
exceed 85 mm. in length. A test board submerged at Bahia, Brasil on May 8, 1945 and
removed on June 2, 1945 contained several specimens of M. striata with the callum com-
pletely developed, the largest specimen measuring 16 mm. in length.

This species was introduced into Sydney Harbour, Australia about ten years ago and is
now becoming very destructive. Life history studies carried on by D. D. Moore’ indi-
cate that at this locality it breeds only during the warmest months of the year, that
the eggs are fertilized in the water and the larvae are free-swimming for about one month.
He states that ‘‘during the winter nearly all the animals of this species change sex to
maleness, the warmer temperature causing a proportion of these to change to female. ”’
This statement, however, needs verification.

Martesia striata Linné, like other callum-building pholads, upon reaching maturity
stops boring and closes off the anterior end of the shell with a calcareous deposit. The
time when this takes place appears to vary greatly with the conditions of substrata and
crowding of the specimens. We have seen minute specimens less than a month old with
the callum fully developed and other large specimens over three months old which were
still active. Working specimens when removed from their burrows are completely help-
less and soon die. However, if the burrow is broken away exposing the posterior portion
of the shell while leaving the anterior portion in the burrow, so that the animal can bring
the shell into action, it will rapidly re-bury itself. A specimen observed burrowed about
22 mm. in 72 hours. This specimen was boring in palmetto piling. Undoubtedly one
working in a harder substance would progress much more slowly.

Martesia striata Linné has been recorded by Pulteney and others from the British Isles,
but these records, like those from Massachusetts, appear to be fortuitous. Undoubtedly
they are based upon specimens taken from wooden ships that had been in the West In-
dies or from driftwood that had been carried by the Gulf Stream.

1D. D. Moore, Jan. 1947, Port of Sydney Journal 1, no. 3, pp. 74-75.

110 JOHNSONIA, No. 34 Martesia

Range. WersTERN ATLANTIC: From Kure Beach, North Carolina south to Florida,
the Gulf of Mexico, the West Indies and south to Rio de Janeiro, Brasil.

EaAsTeERN Paciric: From Sonora, Mexico south to the Gulf of Guayaquil, Peru.

Inpo-Paciric: Hawaiian Islands west to Japan and south through the East Indies to
Brisbane, Australia.

Specimens examined. \WESTERN ATLANTIC: NortH Carouina: Kure Beach (MCZ).
Sourn Caronina: Sullivans Island, Charleston (Charleston Museum). FLortpa: May-
port; Daytona Beach; Fort Pierce (all MCZ); West Palm Beach (USNM); Miami
(A. Merrill); Bahia Honda Key (ANSP); Stock Island, near Key West (USNM);
Estero Island, Fort Myers Beach: Sanibel Island (both MCZ); Boca Grande (ANSP);
Punta Gorda (MCZ; USNM; ANSP); Manatee River, near Bradenton (A. Koto);
Anna Maria (USNM); St. Petersburg (MCZ; ANSP); Gulfport (MCZ); Cedar Keys
(MCZ; USNM); Port St. Joe (A. Merrill); Panama City (MCZ); Pensacola (MCZ;
USNM). Texas: Port Isabel (MCZ). Cuspa: Mariel, Pinar del Rio (Museo Poey);
Banes, Oriente; Guantdinamo, Oriente; La Milpa, Cienfuegos, Las Villas (all MCZ);
Santiago de Cuba, Oriente (USNM). Jamaica: Port Royal; Kingston (both MCZ);
Port Morant; St. Andrews (both USNM). Hispanioua: 'Torbeck; Aquin; Les Cayes;
St. Louis; all Dept. du Sud, Haiti (all USNM). Puerto Rico: San Juan (MCZ;
USNM). Virein Isuanps: St. Croix (USNM; ANSP); St. Thomas (MCZ; USNM;
ANSP: Redpath Museum). Lesser ANTILLES: Macqueripe Bay; Claxtons Bay; Ota-
heite, 5 miles southwest of S. Fernando; all Trinidad (all MCZ). Mexico: Tecolutla,
Vera Cruz (T. Pulley). GuaremMaLa: Puerto Barrios (MCZ). Honpuras: Puerto Cor-
tez(MCZ). Nicaracua: Greytown(USNM). Panama: Chagres; Fort Sherman; Crist-
obal; Coco Solo; Fort Amador (all MCZ). VENEZUELA: Amuay Bay; Puerto Cabello
(both MCZ). Brasii: Bahia, Bahia(MCZ); Mariqui, Bahia de Sepetiba, Rio de Janeiro;
Nictheroy, Rio de Janeiro (both H. Lopes); Praia Boa Viagem, Rio de Janeiro (W. J.
Eyerdam); Cananeia, Sao Paulo (J. P. Carvalho).

Eastern Paciric: Mexico: about 43 mi. S.E. of Altata, Sinaloa (N. Lat. 24°22’;
W. Long. 107°37’); Bahia de Topolobampo (both R. J. Menzies); Guayamas, Sonora
(USNM):; Empalme Beach, Sonora (S. S. Berry); Magdalena Bay, Baja California
(USNM; California Acad. Science). Barra de Cahuacan, Chiapas (Hancock Founda-
tion). NicaraGua: Corinto (California Acad. Science). Costa Rica: Golfito (MCZ);
San Lucas Island, Gulf of Nicoya (USNM). Panama: Puerto Armuelles; Panama,
Canal Zone; Balboa, Canal Zone (all MCZ); Bahia Honda (N. Lat. 70°45’; W. Long.
81°31’) (Hancock Foundation). CotomsBia: Isla del Gallo, Rada de Tumaco (A. A.
Olsson). Ecuapor: Sua (A. A. Olsson). Peru: Tumbes (USNM).

Inpo-Paciric: HawarAn IsLaNnps: Pearl Harbor, Oahu (MCZ; USNM). Japan:
Yawatahama, Ehime, Shikoku (T. Habe); Sasebo, Kyushu (MCZ); Shirahama and
Tanabe Bay, Wakayama, Honshu (both T. Habe). Mariannas IsLanps: Guam (MCZ).
PHILIPPINE IsLANDS: Subic Bay (MCZ; USNM); Mariveles, Bataan (USNM); Cavite,
Manila Bay (MCZ; USNM); off Corregidor Light, Manila Bay: Ragay River, Ragay
Gulf, all Luzon; off Mariquitdaquit Island, Leyte; Dapitan, Mindanao; Malapaya Bay,
Palawan (all USNM). Easr INpies: South of Dodeps Island, Gulf of 'Tomini, Celebes

Western Atlantic JOHNSONIA, No. 34 111

(USNM); Island of Raha, south Celebes; Soengailiat, Banka Island; Rotterdam and
Dapoer, Batavia Bay, Java: Wahai, Ceram; Molucca Islands (all Zool. Mus. Amster-
dam). CuHiNna: off Pratas Island, China Sea (USNM). Srrairs SerrLeEMENts: Johore
Straits (R. D. Purchon). Avsrraia: Brisbane (C. J. Trist).

Martesia (Martesia) fragilis Verri/l and Bush
Plates 65-66

Martesia (Martesiella) fragilis Verrill and Bush 1890, Proceedings United States National Museum 20, no.
1139, p. 777, pl. 79, fig. 10 (in floating wood about 360 miles off Cape Charles, Virginia, Albatross, station
2566).

Pholadidea (Penitella) minuscula Dall 1908, Bulletin Museum Comparative Zodlogy 43, p. 425 (boring in
corky envelope of large seed, Gulf of Panama, Albatross, station 3392).

Martesia (Diploplax) exquisita Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104, no.
11, p. 10, pl. 3, figs. 17-18 (Stony Cove, St. Mary’s Parish, Jamaica).

Martesia (Diploplax) bahamensis Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104, no.
11, p. 11, pl. 3, figs. 15-16 (from a floating nut, South Bight, Andros Island, Bahama Islands).

Distinctive characters. Shell small, not reaching over ? inch in length. Mesoplax broadly
oval and depressed with a peripheral keel and concentric sculpture. The mesoplax as-
sumes the adult form in immature specimens. Metaplax and hypoplax similar to M.
striata Linné.

Description. Shell reaching 19 mm. (about ? inch) in length and 11.5 mm. (about 2
inch) in height, generally pear-shaped and producing a callum in the adult stage. Young
specimens beaked anteriorly and widely gaping: the beaks being truncated at nearly a
right angle, give the shells a teredo-like appearance. Shells rounded and closed posteri-
orly. Valves divided into two distinct portions by a shallow umbonal-ventral sulcus. An-
terior portion sculptured with close-set, concentric, denticulated ridges, and weak to
moderately pronounced radial ribs. Posterior portion sculptured with smooth, rounded
ridges. Umbos prominent and located very near the anterior end of the shell in adult
specimens. In young specimens the umbos are nearly central, but as the posterior slope
elongates, the umbos become more anteriorly located. Umbonal reflections simple, thin,
free anterior to the umbos and broadly recurved forming a funnel-shaped pit below the
reflection. In addition, there is a sickle-shaped flange extending over the umbos which
is closely attached anteriorly but free posteriorly, and raised well above the surface of the
shell. This flange is the attachment area for the anterior adductor muscle. In the adult
the pedal gape is closed by a thin callum which does not extend dorsally between the
beaks. The narrow slit remaining between the two halves of the callum is covered with
a thin but strong periostracum leaving only a minute anterior pore open in the fully adult
specimen. Mesoplax in very young specimens transverse, more or less semicircular in
outline with a shallow U-shaped to deep V-shaped embayment anteriorly. It is divided
longitudinally into two parts by a pronounced groove and is sculptured by concentric
growth lines. The dorsal portion of the mesoplax is produced before the callum is com-
plete so that immature specimens may also occur with nearly complete dorsal plates.
Mesoplax in adult specimens circular to oval in outline, depressed and with a pronounced
concentric sculpture. Metaplax long and narrow, pointed anteriorly and broadening pos-

112 JOHNSONIA, No. 34 Martesia

teriorly to a rounded or truncate posterior margin. Hypoplax long and narrow, pointed
anteriorly, rounded posteriorly and extending from the umbonal-ventral sulcus posteriorly
about two thirds the distance to the posterior margin.

Interior of the shell white and glazed. Umbonal-ventral sulcus expressed internally as
a pronounced ridge. Apophyses long, thin and extending from beneath the umbos an-
teriorly at a moderate angle. Muscle scars well marked, pallial sinus broad and deep,
extending anteriorly in an irregular curve nearly to the umbonal-ventral ridge. Perios-
tracum thin, light straw-yellow in color and persistent.

\We have not seen the soft parts of this species.

Plate 65. Martesia fragilis Verrill and Bush. Fig. 1. Dorsal view of the lectotype of Martesia fragilis showing
the concentrically sculptured mesoplax (8x). Fig. 2. Dorsal view of an adult specimen from Galveston,
Texas, showing the mesoplax and the metaplax (3x). Fig. 3. Paratype of Martesia (Diploplax) bahamensis
Bartsch and Rehder (= M. fragilis Verrill and Bush, young) showing the mesoplax which is essentially
similar to the adult form even though the callum is not produced (123x). Fig. 4. Internal view of an adult
specimen from Galveston, Texas (3x). Fig. 5. Ventral view of an adult specimen from Galveston, Texas
(3x). Fig. 6. Holotype of Martesia ( Diploplax) exquisita Bartsch and Rehder (= M. fragilis Verrill and Bush,
young) (about 43x). Fig. 7. Holotype of Martesia (Diploplax) bahamensis Bartsch and Rehder (= M. frag-
iis Verrill and Bush, young) (about 45x). Fig. 8. Side view of an adult specimen showing the fine sculp-
ture on the anterior portion and the nearly smooth posterior portion. Galveston, Texas (3x). Fig. 9. Dor-

sal view of young specimen to show the mesoplax with its concentric sculpture and slightly concave upper
surface. Sabine, Texas (3x).

Western Atlantic JOHNSONIA, No. 34 113

length * height ratio h:]
19.5 mm. 11.5 mm.

~l

iF Galveston, Texas
ae
1

jy fa) 6.5 6 Matagorda Island, Texas
14.0 8.2 =i Port Aransas, Texas

1225 6.2 2.0 Modesto, Sinaloa, Mexico
1.5 5.0 LD Paratype

6.0 8.5 La Lectotype

* All specimens measured had a complete callum

Types. The following types are all in the United States National Museum. Lectotype,
Martesia fragilis Verrill and Bush, no. 52543, from Albatross, station 2566, about 360
miles off Cape Charles, Virginia (N. Lat. 37°23’; W. Long. 68°08’) in a piece of floating
wood. Holotype of Martesia (Diploplax) exquisita Bartsch and Rehder, no. 573548, from
Stony Cove, St. Mary’s Parish, Jamaica. Holotype of Martesia (Diploplax) bahamensis
Bartsch and Rehder, no. 573549, taken from a floating nut [ T’erminalia catappa Linné *}
collected on the eastern end of South Bight, Andros Island, Bahama Islands. Holotype
of Pholadidea (Penitella) minuscula Dall, no. 122946, from Albatross, station 3392, in the
Gulf of Panama (N. Lat. 7°5’; W. Long. 79°40’) in a nut [Malpighia sp."}.

Remarks. This species is closely related to M. striata Linné and has long been con-
fused with it. Good complete specimens of M. fragilis may be readily differentiated,
however, by the characteristic mesoplax which is depressed, has sharply keeled edges and
pronounced concentric sculpture. In addition, the metaplax of M. fragilis is proportion-
ately shorter and rounded to truncate posteriorly. The young of these two species are
often impossible to differentiate. However, when still immature, WM. fragilis begins the
production of the dorsal portion of the mesoplax and at this stage they may be readily
separated.

Similar to other species in this genus, M. fragilis is rather variable in size and shape,
and lacking its accessory plates could not with certainty be distinguished from M. striata
Linné. Martesia (Diploplax) exquisita Bartsch and Rehder, and Martesia (Diploplax)
bahamensis Bartsch and Rehder appear to be only the young of this species.

We do not understand at present the distribution of this species. The only area where
specimens have been taken in fixed structures (i.e., test boards) is the Texas coast and
here their occurrence is very irregular both as to season and duration. All other records
have come from floating wood or nuts. It would appear that the species is essentially

Plate 66. Mesoplax of Martesia fragilis Verrill and Bush. Fig. 1. Mesoplax of the holotype of Pholadidea
(Penitella) minuscula Dall (= M. fragilis Verrill and Bush). Figs. 2-4. Mesoplax of the paratype of Martesia
Sragilis Verrill and Bush. Fig. 2. Dorsal view showing the concentric growth lines. Fig. 3. Ventral view.
Fig. 4. Side view.

'Dr. Albert F. Hill, Botanical Museum, Harvard University, kindly made the determinations.

114 JOHNSONIA, No. 34 Martesia

pelagic and probably is far more evenly distributed throughout the West Indies than
the few spotted records would indicate.

Range. WesverN ArLantic: In sporadic localities ranging from off Cape Charles,
Virginia, in floating wood, south through the Gulf of Mexico and the West Indies to
Rio Grande do Sul, Brasil.

Eastern Paciric: From Sonora, Mexico south to the Gulf of Panama.

Specimens examined. WESTERN ATLANTIC: Vireinia: Albatross, station 2566 (N.
Lat. 87°23’: W. Long. 68°08’) about 360 miles off Cape Charles (USNM). Fioripa:
Hollywood (ANSP); Oceanus (USNM). Louisiana: Grand Isle (USNM). Texas:
Sabine (MCZ): Galveston (ANSP; T. E. Pulley); Matagorda Island (USNM);: Port
Aransas (J. Hedgpeth); Rockport (MCZ); St. Josephs Island (USN M):; Corpus Christi
(MCZ). Mexico: Tampico, Tamaulipas; Progresso, Yucatan (both ANSP). Bermupa:
(MCZ). Banama IsLanps: eastern end, South Bight, Andros Island (USNM). Ja-
MaIca: Stony Cove, St. Mary’s (USNM). Brasii: Rio Grande do Sul (MCZ; Univ.

of Michigan).

Eastern Paciric: Mexico: Bacochibampo Bay, Sonora; Mendia, Sinaloa; near
Modesto, Sinaloa (all USNM). Panama: Albatross, station 3392, Gulf of Panama (N.
Lat. 7°5’; W.Long. 79°40’) (USNM).

Subgenus Particoma Bartsch and Rehder

Particoma Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104, no. 1, p. 5.

Species in the subgenus Particoma are characterized by having a thick, oval-shaped cal-
lus over the umbos and obliquely to sinuously truncated beaks. The mesoplax is cunei-
form with a central groove from which growth lines radiate. The metaplax and hypoplax
are long, narrow and divided posteriorly.

Subgenotype, Pholas cuneiformis Say, original designation.

Martesia (Particoma) cuneiformis Say
Plates 67-68

Pholas cuneiformis Say 1822, Journal Academy Natural Sciences Philadelphia 2, p. 322 (the southern coast
[United States] ).

Pholas caribaea d’Orbigny 1842 [in] Sagra, Histoire de l’Isle de Cuba, atlas pl. 25, figs. 20-22; d°Orbigny
1853, ibid., text 2, p. 216 (Cuba; Mexico).

Pholas falcata Sowerby [in] Reeve 1872, Conchologia Iconica 18, Pholas, pl. 12, sp. 51 (Hudson’s Bay);
non Wood 1815.

Pholas krebsi “C.B.Adams’ Krebs 1864, The West Indian Marine Shells with some Remarks, Nykjébing,
Denmark, p. 113 [nomen nudum].

Distinctive characters. Shell small, under one inch in length, pear-shaped, widely gap-
ing anteriorly in the young stage and producing a callum when adult. Beaks sinuously
truncate. Mesoplax cuneiform and sculptured with pronounced growth lines radiating
from a median longitudinal groove. Metaplax and hypoplax divided posteriorly.

Western Atlantic JOHNSONIA, No. 34 115

Description. Shell reaching 21 mm. (about £ inch) in length and 11.5 mm. (about 4
inch) in height. Generally more or less pear-shaped in outline and producing a callum in
the adult stage. Young specimens widely gaping anteriorly with obliquely and sinuously
truncated beaks. Shell rounded and closed posteriorly. Valves divided into two distinct
portions by a narrow but well impressed umbonal-ventral sulcus. Anterior portion sculp-
tured with close-set, concentric, denticulated ridges. Radial ribs slightly indicated in
some specimens. Posterior portion sculptured with smooth, rounded, concentric ridges
and faint growth lines. Umbos prominent and in adult specimens located very near the
anterior end of the shell. In very young specimens the umbos are nearly centrally located.
Umbonal reflection small and formed as a thick callus closely appressed over the umbos.
A small funnel-shaped pit is formed below the reflection where it is free anterior to the
umbos. The anterior adductor muscle scar shows as a rather deep oval depression in the
umbonal reflection. In the adult the pedal gape is closed by athin callum which extends
dorsally between the beaks as a narrow margin. The slit remaining between the two
halves of the completed callum is closed by a thin periostracum, leaving only a minute
anterior pore open in the adult specimen. The mesoplax in young specimens (those under
4 mm. in length) is transverse and located in the normal position posterior to the umbos
and beneath the posterior portion of the anterior adductor muscle. However, while the
animal is still immature the mesoplax assumes the adult form and produces the dorsal
portion which is more or less cuneiform in outline. It may range in shape from about
twice as long as wide to nearly square and the median groove may be strongly marked
or barely visible. Sculpture of the mesoplax consists only of pronounced growth lines
radiating from the median groove. Metaplax long and narrow and, in fully adult speci-
mens, is divided posteriorly, extending over the edges of the valves and fusing with

Plate 67. Martesia cuneiformis Say. Fig. 1. Dorsal view of an adult specimen to show the cuneiform mesoplax
and the divided metaplax. Fig. 2. Side view of an adult specimen to show the chitinous *“bulbs’’ at the
posterior end of the valve and the triangular calcareous projection produced by the fusing of the divided
ends of the mesoplax and the metaplax. This condition is only found in old specimens. Fig. 3. Ventral
view of an adult specimen to show the callum and the divided hypoplax. Fig. 4. Dorsal view of a young
specimen to show the mesoplax which is essentially similar to that of the adult. Fig. 5. Ventral view of a
young specimen to show the pedal gape, the apophyses, and the chondrophore with the internal ligament
still in place. Fig. 6. External view of the valve of a young specimen to show the umbonal-ventral sulcus
and the rather coarse sculpture on the posterior slope. Fig. 7. Internal view of the valve of a young speci-
men to show the apophysis and the nearly smooth umbonal area which has only a callus but no flange.

Figs. 1-3. San Juan, Puerto Rico; Figs. 4-7. Kure Beach, North Carolina (all atx),

116 JOHNSONIA, No. 34 Martesia

the divided hypoplax. Hypoplax narrow and extending posteriorly from the base of the
umbonal-ventral ridge. In occasional specimens an incipient siphonoplax is formed. This
appears as a pair of thin, triangular, calcareous projections from the posterior margin of
the valves.

Interior of the shell white and glazed. Umbonal-ventral sulcus expressed internally as
a strong and usually beaded ridge, which in young specimens forms a small condyle at
the ventral margin. In the adult stage the ventral margin of the shell is built out beyond
the condyle. Apophyses long, thin and nearly parallel with the umbonal-ventral ridge.
Muscle sears well marked, the anterior adductor scar being more or less reniform and
covering most of the umbonal reflection. ‘The posterior adductor scar is a long oval. Pal-
lial sinus broad and deep, extending anteriorly beyond the umbonal-ventral ridge. Peri-
ostracum thin, light straw-yellow and persistent.

We have not seen material sufficiently well preserved to permit a description of the
soft parts of this species.

length * height ratio h:]
20.5 mm. 11.5 mm. ier Cumana, Venezuela
19.0 9.0 oF San Juan, Puerto Rico
14.5 9.5 15 Wilmington, North Carolina
7.5 5.5 1.4 im Ss +5
~~ p- ~ x ee ee ee
3.9 SI5S: 1.6

* All specimens measured had a complete callum

Types. The holotype of M. cuneiformis Say is in the Academy of Natural Sciences
Philadelphia, no. 50803. The type locality is here restricted to Charleston, South Caro-
lina, a locality from which Say received considerable material. The type of Pholas caribaea
d’Orbigny is in the British Museum according to Gray 1854; the type locality is here
restricted to Guantanamo, Cuba. The location of the type of Pholas falcata Sowerby is
unknown. It is not in the British Museum according to a letter received from G. L.
Wilkins. The type locality, Hudson Bay, is unquestionably an error.

Remarks. This species, though variable, is very distinctive and can readily be distin-
guished by its cuneiform mesoplax, its small, closely appressed umbonal reflections, and
its divided metaplax and hypoplax. From Diplothyra smithii Tryon, with which it has
been confused, it differs by having a regularly sculptured, cuneiform mesoplax and by
not having the callum extended on either side of the mesoplax. In addition, it is a
wood-borer.

It has been impossible to separate Martesia cuneiformis Say from M. caribaea a’ Orbigny.
These two forms have generally been separated on the basis of the shape of the mesoplax,
but it is possible to obtain specimens exhibiting the complete range of variation from a
short and broad to a long and narrow mesoplax in a single test board. This variation in
shape appears to be related to the rate of growth and the amount of crowding in the speci-
mens. There is also a rather wide range of variation in the number of denticulated ridges
and the size of the denticulations. As in other groups throughout the Pholadidae, these
characters cannot be used safely in distinguishing species because they are readily affected
by the material in which the animal is boring.

Though specimens of MM. cuneiformis Say are recorded from New Jersey and C. W.

Western Atlantic JOHNSONIA, No. 34 117

Johnson reports this species from Connecticut, it seems probable that its occurrence at
these northern localities is fortuitous. The Clapp Laboratories have had test boards all
along the coast from Connecticut to Florida for a number of years but no specimens have
been taken north of Wilmington, North Carolina. It would appear, therefore, that it
is unable to breed in northern waters. The above-mentioned records were probably based
upon specimens carried north in driftwood by the Gulf Stream, or by wooden ships that
had been cruising in the West Indies.

Little is known of the biology of this species. It is often found living in the same situ-
ation as Martesia striata Linné, though it is usually much less abundant. Its range in the
Western Atlantic nearly coincides with that of M. striata, but it has only been taken at
one locality in the Eastern Pacific. It is quite possible that this record may be based upon
a recent introduction into the Pacific by way of the Panama Canal. It is impossible at
present to explain why this species has not become widely distributed as has M. striata
Linné.

From test-board records it would appear that this species breeds in San Juan, Puerto
Rico from September to November and at Bahia, Brasil in May and June. A board sub-
merged May 8, 1945 at Bahia, Brasil had specimens with a fully developed callum on
June 2, 1945. One of these specimens had reached a length of 11.5 mm., indicating
rather rapid growth.

Range. From Cartaret County, North Carolina south through the West Indies and
Central America to Santos, Brasil. Johnson (1934) gives the range as from Connecticut
south, but it would appear that this record was based on specimens that were advectitious
at this locality (see Remarks).

Records. New JERSEY: Holly Beach; Atlantic City (both ANSP). Norra Carouina:
Core Creek Canal, Carteret Co. (Univ. of Michigan); Harbor Island (MCZ): Beaufort
(USNM); Kure Beach; Wilmington; Southport (all MCZ). Sourn Carortina: Myrtle
Beach; Pawleys Island (both Charleston Museum); Isle of Palms; Sullivans Island
(both Charleston Museum; USNM); Charleston Bay (USNM); Folly Beach (MCZ):
Edisto Island (USNM). Grore1a: Fort Screven, Chatham Co. (Charleston Museum);
Sea Island, St. Simon Island (USNM). FLortpa: Mayport (MCZ); St. Augustine
(MCZ: USNM): Daytona Beach; Cape Canaveral (both MCZ); Cocoa Beach (USN M);
Cape Florida(USNM); Key West (Charleston Museum); Fort Myers Beach (ANSP);

Plate 68. Mesoplax of Martesia cuneiformis Say. The above series shows the range in shape of the mesoplax
as expressed by specimens taken from a single test board which was submerged at Mayport, Florida. Fig.
1. Dorsal, ventral and side view of a typical elongate plate of M. cunejformis Say. Fig. 2. An intermediate

shaped plate. Fig. 3. A broad and nearly square plate.

118 JOHNSONIA, No. 34 Diplothyra

Sanibel Island; Captiva Island (both MCZ); Sarasota Bay (USNM); Maximo Point,
St. Petersburg (M. Teare); Cedar Keys (USNM; ANSP; Univ. of Michigan); Port
St. Joe (A. Merrill); Panama City; Pensacola (both MCZ). ALABAMa: Fort Morgan;
Mobile (both USNM). Mississrprr: Pascagoula (USNM). Louisiana: Grand Lake,
Cameron Co. (USNM). Texas: Galveston(MCZ; USNM); Harbor Island, Port Aran-
sas(MCZ). Cusa: Cabanas Bay, Pinar del Rio (Museo Poey); Banes, Oriente; Guan-
tinamo Bay, Oriente (both MCZ). Jamaica: Kingston (MCZ). Hispantouta: Puerto
Plata, Santo Domingo (MCZ); Bezanton, Dépt. de L’Ouest, Haiti; Aquin, Dépt. du
Sud, Haiti (both USNM). Purerro Rico: San Juan; Mayagiiez (both MCZ). VirGin
IsLANDs: St. Thomas(MCZ; ANSP). Panama: Fort Sherman; Cristobal; Coco Solo,
all Canal Zone (all MCZ). VENEzUELA: Amuay Bay; Puerto Cabello; Cumana (all
MCZ). Brasiu: Bahia (MCZ); Santos (USNM).

Eastern Paciric: Panama: Balboa, Canal Zone (MCZ).

Genus Diplothyra T?-yon

Diplothyra Tryon 1862, Proceedings Academy Natural Sciences Philadelphia 14, p. 449; Lamy 1926, Jour-
nal de Conchyliologie 69, p. 194; Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104,
Minos JL jos eye

Shells small, reaching about 15 mm. in length, pear-shaped in outline, divided into
two distinct regions by an umbonal-ventral sulcus and producing a callum in the adult
stage. Shells beaked anteriorly and widely gaping in the young stage, the beaks being
obliquely truncated. Posteriorly the valves are broadly rounded and closed. Umbonal
reflections thin, wide and closely appressed for their entire length. Callum produced dor-
sally between the beaks and extending posteriorly on either side of the mesoplax. Proto-
plax lacking. Mesoplax subquadrate to subtriangulate and extending anteriorly in adult
specimens to cover completely the anterior adductor muscle. Metaplax and hypoplax
pointed anteriorly and forked posteriorly. The forked portions of these plates extend over
the posterior margins of the valves and fuse with one another. Ventral condyles prominent
in young specimens. Apophyses long and thin. Chondrophore and internal ligament
well developed.

Though Diplothyra has been placed by Lamy (1926) and Bartsch and Rehder (1945)
as a subgenus of Martesia the differences expressed in the mesoplax, the umbonal reflec-
tion, the extension of the callum on either side of the mesoplax, the ventral condyles and
the chondrophore seem sufficient to warrant considering this a separate genus. In addi-
tion, species in this genus bore into rock and shells while species of Martesia are wood-
borers. There are only two known species in this genus, one in the Western Atlantic,
the other in the Eastern Pacific.

Genotype, Diplothyra smithi Tryon, monotypic.

Diplothyra smithii 7ryon
Plates 2; 69-71

Diplothyra smithii Tryon 1862, Proc. Academy Natural Sciences Philadelphia 14, p. 450, text figure (Tot-
tenville, Staten Island, New York); Tryon 1863, Proc. Academy Natural Sciences Philadelphia 15, p. 145;
Johnson, C. W. 1905, Nautilus 18, p. 102.

Martesia (Diplothyra) smithii Tryon, Bartsch and Rehder 1945, Smithsonian Miscellaneous Collections 104,
noel, p. 9.

Western Atlantic JOHNSONIA, No. 34 119

Distinctive characters. Shell small, reaching about one half inch in length. Anterior
portion triangular in outline. Callum imbedding the beaks and extending posteriorly on
either side of the mesoplax. Mesoplax of adult specimens composed of a posterior portion
which is concentrically sculptured and an anterior portion which is sculptured only by
irregular wrinkles. Mesoplax and hypoplax forked posteriorly. Usually found in oysters
and coquina rock.

Description. Shell small, reaching 15.5 mm. (about $ inch) in length and 9.5 mm.
(about 4 inch) in height, pear-shaped in outline, white to light brown in color, solid in
structure and producing a callum in the adult stage. Immature specimens beaked and
widely gaping anteriorly; rounded and closed posteriorly. Shell divided into two well
defined areas by an umbonal-ventral sulcus. Anterior portion triangular in outline, sculp-
tured by exceedingly fine, close-set, undulating, concentric ridges and numerous indis-
tinct radial ribs. Posterior portion sculptured only by rather strong growth lines. Umbos
prominent and located near the anterior fourth of the shell in normal specimens. Um-
bonal reflections thin, rather wide and very closely appressed for their entire length. A
pronounced ridge extends longitudinally over the umbos. In the adult the pedal gape is
closed by a strong callum with only a very narrow slit remaining between the two halves.
The slit is covered by a heavy periostracum, leaving only a minute anterior pore open in
fully adult specimens. The callum extends dorsally between the beaks, over the umbonal
reflection and posteriorly on either side of the mesoplax for nearly half its length. Proto-
plax lacking. Mesoplax in young specimens more or less triangular in outline and marked
with concentric growth lines. The dorsal portion of the mesoplax gradually extends for-
ward as the shell matures and, at the time the callum is produced, an unsculptured por-
tion is added to the mesoplax which extends anteriorly between the dorsal extensions of
the callum. Metaplax pointed and bent downward anteriorly, fitting against and under
the mesoplax. It is divided posteriorly and the two halves extend over the posterior mar-
gins of the valves where they fuse with the two arms of the divided hypoplax. Hypoplax
small and extending anteriorly to the umbonal-ventral sulcus. Siphonoplax lacking.

Interior of shell white and usually glazed. Umbonal-ventral sulcus evident internally
as a low beaded ridge. Muscle scars large and well marked. Pallial sinus broad and deep,
extending anteriorly beyond the umbonal-ventral ridge. Apophyses rather long, thin,

Plate 69. Diplothyra smithii Tryon. Fig. 1. Staten Island, New York, Lectotype. Fig. 2. Myrtle Beach,
South Carolina. Figs. 3-5. Bohicket Creek, Edisto Island, South Carolina. Fig. 3. Dorsal view showing
the extension of the callum on either side of the mesoplax. Fig. 4. Ventral view showing the divided hy-
poplax. Fig. 5. Dorsal view showing the divided metaplax (all adults, all 34x),

120 JOHNSONIA, No. 34 Diplothyra

narrow and fragile, extending from beneath the umbos anteriorly at a moderate angle.
A pronounced chondrophore is present on the left valve.

Siphons white, smooth and short, extending only slightly beyond the end of the shell
and capable of complete retraction within the shell. Foot and mantle white. Foot in
young specimens large, nearly circular in outline and truncate. The foot atrophies in
the adult.

length height ratio h:]

14.5 mm. 8.5mm. Av St. Augustine, Florida

13.0 ORD 1.4 Daytona Beach, Florida

12.5 7.5 1.6 ‘- ne ag

10.0 7.0 1.4 Paratype, Staten Island, New York

7.0 4.5 1 Daytona Beach, Florida

Types. The lectotype of Diplothyra smithu Tryon, here selected, is in the Academy
of Natural Sciences Philadelphia, no. 51061 from Staten Island, New York. Paratypes,
Museum of Comparative Zodlogy, no. 124273 and 124274 from Tottenville, Staten Is-

land, New York.

Remarks. This is a very distinctive species and is not closely related to any of the small
borers in the Western Atlantic. It is nearest to D. curta Sowerby of the Eastern Pacific,
from which it differs in its divided mesoplax which in the adult stage is composed of
an early concentrically-sculptured portion and a later portion which is sculptured only
by irregular wrinkles. See also remarks under D. curta Sowerby.

Diplothyra smithu Tryon is the species usually found boring into oysters. It has, how-
ever, been taken from Fasciolaria gigantea Kiener, and may be common in coquina rock.
We have a single record of D. smithi Tryon boring into floating wood. This is a most
unusual habitat for this species and should be confirmed.

Though D. smithi Tryon may on oceasion be very abundant in specimens of oysters,

Plate 70. Diplothyra smithii Tryon. Figs. 1-3. Young specimens from Daytona Beach, Florida. Fig. 1. Ven-
tral view of opposed valves showing the ventral condyles, the apophyses, and the chondrophore with the
ligament still in place. Fig. 2. Internal view showing the long narrow apophyses and the muscle scars.
Fig. 3. External view showing the triangularly shaped anterior portion (all 43x).

Western Atlantic JOHNSONIA, No. 34 121

they do not appear to injure them sufficiently to be of economic importance to the oyster
fisheries.

Range. From Provincetown, Massachusetts south to Daytona Beach and Sanibel Is-
land, Florida, and west to Texas.

Specimens examined. MassacuusEtrts: Provincetown (MCZ). New York: Totten-
ville, Staten Island (MCZ; USNM; ANSP). Pennsytvania: Philadelphia (USNM).
MaryLanp: Crisfield (USN M); Chesapeake Bay (USNM; MCZ). Vireinia: Norfolk
(MCZ). Norrn Carouina: Morehead City; Beaufort (both USNM). Sourn Caro-
LINA: Bohicket Creek, Edisto Island; Sullivans Island; Myrtle Beach; Pawleys Island;
Magnolia Beach (all Charleston Museum); Isle of Palms; Charleston (both USNM:;:
Charleston Museum). FLoripa: St. Augustine; Lake Worth (both USNM; MCZ);
Daytona Beach (MCZ): Sanibel Island (A. S. Merrill); Tampa; Apalachicola (both
USNM). Lovistana: Barataria Bay; Dutchmans Gap, St. Bernardo Parish (both
USNM): Grand Isle (USNM; MCZ). Texas: Port Lavaca; Matagorda Bay; Keller
Bay, Calhoun Co. (all USNM); Port Aransas (MCZ).

Plate 71. Mesoplax of Diplothyra smithii Tryon. Fig. 1. Side view of the mesoplax of an average adult speci-
men. Fig. 2. Ventral view of the same specimen. Fig. 3. Dorsal view of the same specimen showing the
concentrically sculptured area of the young specimen and the irregular unsculptured area produced at the
time the callum is formed. Figs. 4-5. Dorsal views of the mesoplax from other specimens to show the range
in shape, size and proportion of the two areas. Fig. 6. Side view of the mesoplax of a typical young speci-
men. Fig. 7. Ventral view of an unusually wide mesoplax of a young specimen. Fig. 8. Dorsal view of the
same specimen as shown in Fig. 6. Fig. 9. Dorsal view of the same specimen as shown in Fig. 7. All from
specimens taken at Lake Worth, Florida.

Diplothyra curta Sowerby
Plate 72
Pholas curta Sowerby 1834, Proceedings Zoological Society London, p. 71 (Isle of Lions, Province of Vera-

gua, Columbia [Isla Leones, Golfo de Montijo, Prov. of Veraguas, Panama] in soft stone); Sowerby 1849,
Thesaurus Conchyliorum 2, pt. 10, p. 494, pl. 104, figs. 33-34, pl. 108, fig. 105.

Distinctive characters. Shell small, reaching about one half inch in length. Callum
imbedding the beaks and extending posteriorly on either side of the mesoplax. Mesoplax
of adult specimens subquadrate in outline, uniform throughout and nearly smooth. Meta-
plax and hypoplax forked posteriorly. Found normally in shells and soft stone,

122 JOHNSONIA, No. 34 Diplothyra

Description. Shell small, reaching about 15 mm. (about 4 inch) in length, pear-shaped
and producing a callum in the adult stage. Immature specimens beaked and widely gap-
ing anteriorly: rounded and closed posteriorly. Valves divided into two well defined
areas by an umbonal-ventral sulcus. Anterior portion triangular in outline and sculptured
by close-set imbricated ridges and radial ribs. Posterior portion sculptured only by faint
growth lines. Umbonal reflections broad and closely appressed. In the adult the pedal
gape is closed by a callum which extends dorsally between the beaks, usually imbedding
them, and posteriorly on either side of the mesoplax for about half its length. Protoplax
lacking. Mesoplax subquadrate in outline, with faint concentric sculpture and with a
small point in the center of the anterior and posterior margins. Metaplax and hypoplax
forked posteriorly, the forked portions extending over the posterior margins of the valves
and fusing with each other in fully adult specimens. Interior of the shell similar to that
of D. smithu Tryon.

length height ratio h:]
15.5 mm. 8.5 mm. 1.8 Lectotype
11.0 S55 133 Punta San Felipe, Baja California

Types. The types of Pholas curta Sowerby may be in the British Museum. The type
locality as given by Sowerby is ‘‘ad littora Columbiae occidentalis. From the Isle of Lions,
Province of Veragua [Isla Leones, Golfo de Montijo, Prov. Veraguas, Panama], in soft
stone at low water,’” Hugh Cuming collector. The label on the cotype specimens in the
British Museum gives Bay of Caraccas, West Colombia { Bahia de Caraéquez, Ecuador},

| ae hl

3 4 5

Plate 72. Figs. 1-2. Original sketches by P. P. Carpenter of the type specimens of Martesia intercalata Car-
penter (= Penitella conradi Valenciennes). Figs. 3-5. Diplothyra curta Sowerby, Cotypes (2x). Fig. 3. Dor-
sal view showing the subquadrate mesoplax and the forked metaplax. Fig. 4. Dorsal view of an adult
specimen lacking the mesoplax but showing the dorsal extension of the callum which extends on either side
of the mesoplax. Fig. 5. Ventral view of an adult specimen showing the large callum and the forked hypoplax.

All photographs received through the kindness of Mr. G. L. Wilkins, British Museum (Natural History).

Eastern Pacific JOHNSONIA, No. 34 123

Hugh Cuming collector. We figure these specimens, photographs of which were received
through the kindness of G. L. Wilkins of the British Museum.

Remarks. Diplothyra curta Sowerby is very close in its relationship to D. smithi Tryon
and when more material is available for study these two may prove to be only subspeci-
fically different. The nearly smooth uniform mesoplax of curta is the only character
which distinguishes the two species.

This is an exceedingly rare species and most published references to it seem to have
been based upon malformed specimens of Martesia striata Linné. Nothing is known of
the biology of this species, though it is probably similar to that for D. smithi. It is found
in the intertidal zone and out to depths of at least 10 fathoms.

Range. From the mouth of the Colorado River, Gulf of California south to Libertad,
Ecuador.

Specimens examined. Mexico: Colorado River, Gulf of California (USNM); Punto
San Felipe, Baja California (S. S. Berry). Panama: Balboa, Canal Zone (USNM).
Ecuapor: off Libertad (S. Lat. 2°07'30’; W. Long. 80°56/30”), in 10 fathoms (Han-
cock Foundation).

Genus Parapholas Conrad

Parapholas Conrad 1848, Proceedings Academy Natural Sciences Philadelphia 4, p. 121.

Shell moderate to rather large in size, beaked and gaping anteriorly in the young stage
and producing a callum when adult. Shells broadly rounded to acuminate posteriorly
and closed. Accessory plates consisting of a mesoplax, metaplax and hypoplax. Valves
divided into three distinct regions, the anterior beaked portion being separated from the
dise by an umbonal-ventral sulcus. The posterior slope, which is sculptured with a series
of overlapping chitinous plates, is separated from the disc by a slight groove or ridge ex-
tending from the umbos to the posterior ventral margin. Siphonoplax lacking, being re-
placed by a tube or ‘‘chimney”’’ composed of fine particles which have been ejected from
the siphons and then cemented together.

The genus Parapholas is distributed throughout the temperate and tropical portions
of the Eastern Pacific and the Indo-Pacific, with one species, Parapholas branchiata
Gould occurring in West Africa.

Genotype, Pholas californica Conrad, monotypic.

Key to the American species of Parapholas

1. Adult shells generally under 24 inches in length. Mesoplax enlarged,
extending anteriorly over the beaks .... . =. . 1... s.«.s.. &
Adult generally over 24 inches in length. Mesoplax not enlarged, ex-
tending about one half the distance to the beaks . . P. californica

2. Mesoplax truncate posteriorly. Posterior margin of the shell acuminate.
Chitinous plates on the posterior slope angled . . . . . P. acuminata
Mesoplax lobed posteriorly, extending back on either side of the meta-
plax. Posterior margin of the shell narrowly rounded, chitinous plates
on the posterior slope rounded ............ WF. calva

124 JOHNSONIA, No. 34 Parapholas

Parapholas californica Conrad
Plates 73-77

Pholas californica Conrad 1837, Journal Academy Natural Sciences Philadelphia 7, p. 236, pl. 18, figs. 5-6
(San Diego and Santa Barbara, California in soft rocks).

Pholas janellii Deshayes 1839, Revue Zoologique par la Société Cuviérienne 2, p. 357; Deshayes 1840,
Magasin de Zoologie Guérin-Méneville (2) 2, Mollusques, pls. 14-16 and text (shore of California).

Parapholas californica Conrad 1848, Proceedings Academy Natural Sciences Philadelphia 4, p. 121; Fitch,
J. E. 1953, State of California, Dept. of Fish and Game, Fish Bulletin, no. 90, p. 96.

Distinctive characters. Adult shell usually over 24 inches in length. Mesoplax extend-
ing about one half the distance to the beaks. Metaplax pointed posteriorly. Hypoplax
pointed anteriorly, broadly rounded posteriorly. Chitinous plates on the posterior slope
rounded.

Description. Shell reaching 149 mm. (about 5# inches) in length, oval in outline, solid
in structure and producing a callum in the adult stage. [Immature specimens beaked and
gaping widely anteriorly. Shell tapering posteriorly to a broadly rounded or truncate
posterior margin. Valves divided into three well defined regions. Anterior beaked por-
tion sculptured with very close-set, upturned, undulating, concentric ridges and weak
radial ribs. Anterior portion separated from the median area or disc by a pronounced
umbonal-ventral sulcus. Disc sculptured only by pronounced growth lines. Posterior
slope separated from the dise by a ridge extending from the umbo to the posterior ven-
tral margin. Posterior slope, on a shell devoid of periostracum, sculptured with strong

Plate 73. Parapholas californica Conrad. White Point, San Pedro, California (about 2 natural size). Fig. 1.
Ventral view of young specimen showing the pedal gape, the characteristic foot and the overlapping chiti-
nous plates on the posterior slope. Fig. 2. Lateral view of an adult showing the callum, the metaplax and
the hypoplax.

Photographs received through the kindness of J. E. Fitch of the California Fisheries Laboratory, Termi-
nal Island.

Eastern Pacific JOHNSONIA, No. 34 125

growth lines. In fresh and particularly young specimens this portion of the shell is cov-
ered by a series of rounded, overlapping, chitinous plates. Umbonal reflection simple,
narrow and raised slightly above the surface of the umbo. In adult specimens the large
pedal gape is closed by a heavy callum, only a very narrow slit remaining and this is cov-
ered by heavy periostracum except for the small anterior pore. Callum slightly ridged
transversely, indicating the forward extension of the radial ribs. Protoplax lacking, being
replaced by the dorsal extension of the callum and the anterior extension of the enlarged
mesoplax of the adult. Line of contact between the callum and the mesoplax barely vis-
ible in most living specimens and often with an apparent fusion of the two in older speci-
mens. Mesoplax in young specimens thin, small and more or less U-shaped. In the adult,
the dorsal portion of the mesoplax becomes enlarged and produced anteriorly: it is
broadly oval in outline and divided longitudinally by a deep sulcus. In adult specimens
the dorsal margins of the valves posterior to the umbos are reflected and recurved, nearly
enclosing the posterior adductor muscle. ‘The space between the reflections is covered by
a long, narrow metaplax. It is truncate and sharply bent downward anteriorly, fitting
closely against the posterior margin of the mesoplax. Posteriorly the metaplax tapers to
a rounded point. The narrow ventral gape is covered by a long, narrow plate, the hypo-
plax, which is pointed anteriorly and rounded posteriorly. Periostracum ranging in color
from a dark golden-brown to nearly black. It is rather heavy on the disc, and on the
posterior slope it is produced as a series of overlapping plates.

Plate 74. Parapholas californica Conrad. San Diego, California. Fig. 1. Ventral view showing the hypoplax
in place. Fig. 2. The chimney. Fig. 3. Dorsal view showing the metaplax and mesoplax in place and the
dividing line between the mesoplax and the dorsal extension of the callum (all slightly reduced ).

126 JOHNSONIA, No. 34 Parapholas

No true siphonoplax is produced, but instead, there is a characteristic tube or ‘‘chim-
ney” composed of the finely-worked material which is ejected by the siphons and cemented
together to form a thick tube which fits over the posterior end of the shell and into which
a portion of the chitinous plates of the posterior slope may extend.

Plate 75. Parapholas californica Conrad. Santa Cruz, California. Internal view of valve showing the apophy-
sis, the dorsal extension of the callum, the muscle scars and the pallial sinus, and the cavity beneath the

metaplax (14x).

Interior of shell white and glazed. Muscle scars elongate and clearly marked especially
in older specimens. Umbonal-ventral sulcus expressed internally as a strong ridge which
forms a condyle at the ventral margin. Patlial sinus nearly as broad as the shell is high
and extending anteriorly nearly to the ventral condyle. Apophyses rather large, broad,
and spooned at the ventral margin, extending from beneath the umbo anteriorly at a
rather sharp angle.

Siphons not extending more than one and one half times the length of the shell and
capable of complete retraction within the shell. They are white to light ivory for most
of the length, smooth and devoid of periostracum. Incurrent siphon nearly three times

Plate 76. Parapholas californica Conrad. Del Monte Beach, Monterey Bay, California. Young working speci-
mens showing the chitinous plates on the posterior slope, the apophyses, the umbonal-ventral ridge and

the pedal gape.

Eastern Pacifiic JOHNSONIA, No. 34 127

the diameter of the excurrent siphon and surrounded at the opening by numerous branched
cirri. Excurrent siphon nearly smooth. A second ring of cirri surround the combined
siphons just a short distance anterior to the opening. The area between this and the open-
ings is a dark red-gray to red-brown in color and is covered with numerous white papillae.
When extended, the siphons usually appear gray in color. Foot in young specimens
more or less elliptical in outline, rounded anteriorly, pointed posteriorly and truncate.
Foot and mantle white to light ivory in color. The foot atrophies in the adult and the
mantle is completely closed except for a minute anterior pore.

length * height ratio h:]

149.0 mm. 74.5 mm. 2.0 Monterey, California
121.0 67.0 1.7 ¥ y.
115.0 49.5 2.3 San Diego, California
60.5 16.8 3.5 7 i a

* All specimens measured had a complete callum

Plate 77. Accessory plates of Parapholas californica Conrad. Fig. 1. Dorsal view of the metaplax to show the
truncated anterior end and the pointed posterior end. Fig. 2. Ventral view of the metaplax. Fig. 3. Side
view of the metaplax to show the configuration of the anterior end where it bends sharply downward to fit
against the posterior end of the mesoplax. Fig. 4. Internal (dorsal) view of the hypoplax to show the
pointed anterior end, the rounded point of the posterior end and the cavity into which the muscular mar-
gin of the mantle fits. Fig. 5. External (ventral) view of the hypoplax. Fig. 6. Dorsal view of the meso-
plax to show its nearly circular outline and central division. Fig. 7. Ventral view of the mesoplax to show
the cavity into which the anterior adductor muscle fits. Fig. 8. Side view of the mesoplax. Fig. 9. Dorsal
view of the mesoplax of a young specimen. Fig. 10. Ventral view of the mesoplax of a young specimen.

Figs. 1-8. From specimens collected at San Francisco, California. Figs. 9-10. From specimens collected
at White Point, San Pedro, California.

128 JOHNSONIA, No. 34 Parapholas

Types. The location of the type specimens of Pholas californica Conrad is unknown,
the type locality is San Diego, California. The type of Pholas janellii Deshayes is possi-
bly in the British Museum, the type locality is the shores of California.

Remarks. Parapholas californica Conrad is a very distinctive species and is readily dis-
tinguished from all other species in this genus by its much larger size, its divided meso-
plax which may be fused with the dorsal extension of the callum, and its rounded to
truncate posterior margin. See also remarks under P. acuminata Sowerby and P. calva
Sowerby.

Parapholas californica Conrad, along with Penitella penita Conrad and Chaceia ovoidea
Gould, is found boring into clay, shale and soft friable stone, but is never a deep borer.
Mr. John E. Fitch has written that the deepest he has ever had to dig for a P. califor-
nica Conrad was ten inches and that a four-inch specimen would seldom be more than
eight inches down. ‘This species is found wherever proper ecological conditions exist from
the intertidal zone out to depths of 20 to 30 feet.

Range. Bodega Lagoon, Sonoma County, California south at least as far as Bahia San
Bartolomé, Baja California. Dall’s record for Coos Bay, Oregon (Nautilus 1897 11, p.
66) is probably in error.

Specimens examined, CALIFORNIA: Bodega Lagoon, Sonoma County (Hancock Foun-
dation); Santa Cruz (MCZ); Monterey (MCZ; USNM; ANSP; J. E. Fitch); Del
Monte Beach (MCZ; E. P. Chace); Santa Barbara (USNM); San Pedro (ANSP);
White Point, San Pedro (E. P. Chace; USNM); Anaheim Bay (MCZ); Portuguese
Bend, Orange County (Hancock Foundation); La Jolla (MCZ; Charleston Museum);
San Diego(MCZ: USNM: ANSP: Hancock Foundation). Mexico: Bahia San Bar-
tolomé, Baja California (J. E. Fitch).

Parapholas acuminata Sowerby
Plates 78-80
Pholas acuminata Sowerby 1834, Proceedings Zoological Society London, p. 70; Sowerby 1849, Thesaurus
Conchyliorum 2, pt. 10, p. 492, pl. 105, figs. 48-50 (Panama); non Pholas acuminata Conrad 1845; Fischer
1858, Journal de Conchyliologie 7, pp. 169-177, pl. 6, figs. 1-11.
Parapholas acuminata Sowerby, Carpenter 1855, Catalogue of the Reigen Collection of Mazatlan Mollusca,
p. 12; Lamy 1926, Journal de Conchyliologie 69, p. 163.

Distinctive characters. Shell about 24+ inches long. Mesoplax greatly enlarged, com-
pletely covering the beaks anteriorly; truncate posteriorly. Chitinous plates on the pos-
terior slope angled. Metaplax and hypoplax forked posteriorly.

Description. Shell reaching 61 mm. (about 24 inches) in length and 29 mm. (about 14
inches) in height, pear-shaped, heavy, solid in structure and producing a callum in the
adult stage. Immature specimens beaked anteriorly and widely gaping; tapering pos-
teriorly to a narrowly rounded and often pointed posterior margin. Shell divided into
three well defined areas. Anterior portion sculptured with extremely close-set, low, up-
turned, concentric ridges and numerous radial ribs. Ribs indicated mainly by the radial
arrangement of the rows of undulations. Anterior slope separated from the dise by a nar-

Eastern Pacific JOHNSONIA, No. 34 129

row umbonal-ventral sulcus. Disc sculptured by growth lines and separated from the
posterior slope by a slight groove extending from the umbo to the posterior ventral mar-
gin. Posterior slope sculptured by growth lines only on a shell devoid of periostracum.
On fresh specimens it is covered by a series of angled overlapping chitinous plates. Um-
bonal reflections in young specimens narrow, rather thick and free for the entire length.
Dorsal margin of valves posterior to the umbos, only slightly reflected and recurved.
Pedal gape closed in the adult stage by a rather heavy callum with a very narrow slit re-
maining between the two halves; the slit is covered with a heavy periostracum, leaving
only a minute anterior pore in the fully adult specimen. Callum sculptured with faint
growth lines and parallel ridges which are the extensions of the radial ribs. The callum
does not extend dorsally between the beaks. Mesoplax in the adult greatly enlarged and
extended anteriorly, completely covering the anterior adductor muscle and the beaks.
It is broadly oval in outline, terminating anteriorly in a small point which fits in between
the two halves of the callum. The mesoplax of the young specimen is unknown. Meta-
plax long and narrow; rounded and sharply bent downward anteriorly so that it abuts
closely against the mesoplax; tapering and forked posteriorly. Hypoplax pointed anteri-
orly and forked posteriorly. Siphonoplax lacking. Chimney extending down over the
surface of the shell anteriorly as far as the mesoplax. Periostracum dark brown to almost
black in color, heavy on the dise and produced as overlapping angled plates on the pos-
terior slope.

Interior of shell white to grayish-green in color and usually glazed. Umbonal-ventral
sulcus evident internally as a strong ridge which, in young specimens, extends beyond
the ventral margin of the valve to form a condyle. In the adult, the shell is built out
beyond the end of the umbonal-ventral ridge and the condyle. Posterior adductor mus-
cle scar elongate and well marked. Pallial sinus rather small, extending anteriorly about
one half the distance to the umbonal-ventral ridge.

The soft parts of this species were unavailable for study, but according to Fischer

¢
|
}

Plate 78. Parapholas acuminata Sowerby. Figs. 1-2. Panama Bay, one mile off the canal entrance in 10 fath-
oms. Fig. 1. Dorsal view showing the mesoplax, which is truncate posteriorly, and the divided metaplax.
Fig. 2. Side view showing the angled plates on the posterior slope and the extension of the mesoplax an-
teriorly covering the beaks. Fig. 3. Ventral view showing the hypoplax which is pointed anteriorly and
divided posteriorly (all natural size).

130 JOHNSONIA, No. 34 Parapholas

(1858) the siphons are short, capable of complete retraction within the shell, and the open-
ings are surrounded with triangular cirri. The mantle is similar in color to the mantle
of Mytilus.

length * height ratio h:1
61.0 mm. 29.0 mm 2.10 Panama Bay, off the canal
5253 22.0 2.37 Panama
44.5 20.0 2220) Mazatlan, Mexico
34.5 16.9 2.04 Panama Bay
28.0 14.5 1.93 La Libertad, Ecuador

* All specimens measured had a complete callum

Types. The type of Pholas acuminata Sowerby is in the British Museum according to
a letter received from G. L. Wilkins. The type locality is Panama, Hugh Cuming
collector.

Remarks. Parapholas acuminata Sowerby is probably most closely related to P. calva
Sowerby and has often been synonymized with that species. It differs, however, in being
much more acuminate posteriorly, having the chitinous plates on the posterior slope
angled rather than rounded, and by having the mesoplax truncate rather than lobed pos-
teriorly. It is also close in its relationship to Parapholas californica Conrad from which
it differs by being much smaller in size, by being sharply pointed posteriorly and having
a much larger mesoplax, with the metaplax and hypoplax being forked posteriorly.

Parapholas acuminata Sowerby is apparently a very rare species to judge from the few
specimens available for study. It bores into limestone and probably other soft stone and
clay. The burrow is lined with agglutinized particles from its borings so that a ‘‘chim-
ney’’ is produced which extends down over the surface of the shell at least as far as the
mesoplax. It is found living in depths of from two to ten or more fathoms.

Plate 79. Parapholas acuminata Sowerby. Panama. Fig. 1. Inner view of an adult shell showing the elongate
posterior muscle scar. Fig. 2. External view of young specimen showing the high umbonal reflection. Fig.
3. Internal view of young specimen showing muscle scars, pallial sinus and apophysis. Fig. 4. Chimney
(the circular hole in the side of the chimney was made by another borer) (all about 13x).

Eastern Pacific JOHNSONIA, No. 34 131

Range. From San Martin Island, near San Quintin, Baja California south at least as
far as Bocapan, Peru.

Specimens examined. Mexico: San Martin Island, near San Quintin in 2 to 3 fathoms
(USNM); Mazatlan (MCZ; S.S. Berry). Panama: (AMNH; MCZ; USNM); Pan-
ama Bay, one mile off canal entrance in 10 fathoms (Stanford Museum: E. P. Chace).
Ga LapaGos IsLanpDs: Chatham Island (USNM). Ecuapor: Manta (USNM:;: A. A.
Olsson); off La Libertad (S. Lat. 2°12/15’: W. Long, 80°53’40”) in 4 fathoms (Han-
cock Foundation); Crucita; San Pedro, near Manglaralto; Santa Elena Peninsula; Sali-
nas (al] A. A. Olsson). Peru: Bocapaén (A. A. Olsson).

Plate 80. Accessory plates of Parapholas acuminata Sowerby. Fig. 1. Hypoplax showing the forked posterior
end and the pointed anterior end. Fig. 2. Dorsal view of the metaplax showing the broad anterior end
which bends sharply downward and comes to a point. Fig. 3. Side view of the metaplax showing the an-
terior end sharply bent downward to fit against the posterior end of the mesoplax. Fig. 4. Ventral view of
the mesoplax of an adult specimen. Fig. 5. Dorsal view of the mesoplax showing the rounded posterior
margin and the central point on the anterior margin. Fig. 6. Side view of the mesoplax.

Figs. 1-3. From Panama Bay, Panama. Figs. 4-6. From Mazatlan, Mexico.

Parapholas calva Sowerby
Plate 81

Pholas calva ‘Gray’ Sowerby 1834, Proceedings Zoological Society London, p. 69 (Perico Island, Panama
Bay); Sowerby 1849, Thesaurus Conchyliorum 2, p. 493, pl. 105, figs. 51-53; Sowerby 1849 [1850], Pro-
ceedings Zoological Society London, p. 162, pl. 5, fig. 4.

Pholas calva var. nana Sowerby 1834, Proceedings Zoological Society London, p. 70 (Panama).

Parapholas bisuleata Conrad 1849, Proceedings Academy Natural Sciences Philadelphia 4, p. 156 (Lower
California and Peru); Conrad 1850, Journal Academy Natural Sciences Philadelphia 1, pt. 4, p. 279, pl. 39,
fig. 5 [not fig. 4 as given in the text].

Martesia calva Sowerby, Gray 1851, Annals and Magazine Natural History (2) 8, p. 383.

Parapholas calva Sowerby, Carpenter 1855, Catalogue of the Reigen Collection of Mazatlan Mollusca, p. 9;
Hertlein and Strong 1950, Zoologica 35, p. 249.

Distinctive characters. Shell about 1? inches in length. Mesoplax greatly enlarged,
completely covering the beaks anteriorly and extending posteriorly in two lobes on either
side of the metaplax. Metaplax and hypoplax forked posteriorly. Chitinous plates on
the posterior slope rounded.

132 JOHNSONIA, No. 34 Parapholas

Description. Shell reaching 44 mm. (about 1} inches) in length and 25 mm. (1 inch)
in height, white to light brown in color, pear-shaped in outline, rather light in structure
and producing acallum in the adult stage. Immature specimens beaked and widely gap-
ing anteriorly and tapering posteriorly to a narrowly rounded posterior margin. Shell
divided into three well defined areas. Anterior portion rather small, sculptured by close-
set, low, upturned concentric ridges and weak radial ribs. Anterior slope separated from
the disc by a narrow and lightly impressed umbonal-ventral sulcus. Dise sculptured by
close-set growth lines and separated from the posterior slope by a slight groove extend-
ing from the umbos to the posterior ventral margin. Posterior slope covered with a series
of rounded, overlapping chitinous plates. In the adult the pedal gape is closed by a thin
but strong callum, only a narrow slit remaining when the callum is complete. Callum
sculptured with very fine growth lines and faint parallel ridges which are the extensions
of the radial ribs. The callum does not extend dorsally between the beaks. Mesoplax
greatly enlarged in the adult, extending anteriorly to cover the beaks completely. It is
also produced posteriorly into two lobes which extend backward on either side of the
metaplax. The mesoplax of young specimens is unknown. Metaplax long and narrow,
pointed anteriorly, tapering and divided posteriorly. Hypoplax pointed anteriorly, divided
posteriorly. Siphonoplax lacking. Chimney, according to the figure given by Sowerby
(1849), extending anteriorly well over the surface of the shell at least as far as the
umbonal-ventral ridge. Periostracum dark brown in color, heavy on the disc, and _pro-
duced as overlapping plates on the posterior slope. |

It is impossible to describe the interior of the shell as the few specimens available were
intact and to remove the accessory plates and separate the valves seemed unwise. Noth-
ing is known of the soft parts of this species.

length height ratio h:]

49.5 mm. 24.5 2.0 Manta Bay, Ecuador
48.0 24.0 2.0 eran -
44.0 2520 eer Mazatlan, Mexico

Types. The holotype of Parapholas calva Sowerby is in the British Museum according
to a letter received from G.L. Wilkins; the type locality is Perico Island, Canal Zone,
Panama Bay. The location of the holotype of Parapholas bisulecata Conrad is unknown.

Remarks. Parapholas calva Sowerby is apparently a very rare species to judge by the
very few published records and the lack of specimens in collections. It is close in its re-
lationship to P. acuminata Sowerby, but may be distinguished from this species by its
much larger and lobed mesoplax, and its rounded, rather than angled posterior margin.
It is also closely related to Parapholas branchiata Gould, a West African species which
has a similarly lobed dorsal plate (see Nicklés, 1953, Institut Royal des Sciences Naturelles
de Belgique 39, no. 13, p. 10, pl. 1, figs. 10-13).

In the original description of this species Sowerby states that the specimens were taken
from Spondylus from the Isle of Perico, Bay of Panama in 12 fathoms and the young
specimens were taken from hard stone at low water. In the Thesaurus Conchyliorum,
however, he does not mention the Spondylus and though he does not correct this state-
ment, it seems probable that it was in error; at least this would be a most unusual sub-
stratum for a Parapholas. No other species in this genus bores into other shells and the

Western Atlantic JOHNSONIA, No. 34 133

specimens Sowerby figures are much too large to be shell-borers. Carpenter (1855, loc.
cit.) was probably dealing with two species under this name, but without seeing the
specimens with which he was working it is impossible to say definitely.

Range. From Ensenada de San Francisco, Guaymas, Sonora, Mexico to Manta Bay,
Ecuador.

Specimens examined. Mexico: Ensenada de San Francisco, Guaymas, Sonora (Han-
cock Foundation): Mazatlan (MCZ). Ecuapor: Manta Bay in 1 fathom (Hancock
Foundation).

Plate 81. Parapholas calva Sowerby. Mazatlan, Mexico. Fig. 1. Side view showing the rounded plates on the
posterior slope. Fig. 2. Dorsal view showing the lobed mesoplax (about 23x).

Subfamily Jouannetiinae
Genus Jouannetia des Moulins

Jouannetia des Moulins 1828, Bulletin d’Histoire Naturelle Société Linnéenne, Bordeaux 2, p. 244 (geno-

type, Jouannetia semicaudata des Moulins, monotypic).
Touannetia Gray 1840, Synopsis Contents British Museum, ed. 42, p. 154 (emendation for Jouannetia).

Adult shell white to dirty gray-brown in color, globose, inequivalve, with a large pro-
truding callum and no apophyses. In the young stage the shell is equivalved, beaked,
widely gaping anteriorly and closed posteriorly. Anterior portion sculptured with numer-
ous close-set, imbricated ridges and usually weak radial ribs which are indicated by the
radial arrangement of the imbrications. Umbos prominent and in young specimens more

134 JOHNSONIA, No. 34 Jouannetia

or less centrally located. Posterior slope sculptured with concentric growth lines and thin
elevated ridges. In the adult the right valve elongates, producing a siphonoplax which
may be pectinate or smooth. Callum greatly produced giving the adult shell a globose
appearance. On the right valve it is normal, while on the left it is greatly enlarged and
overlaps the right valve. The callum extends dorsally between the beaks and that of the
left valve forms the covering for the anterior adductor muscle. Interior of the shell white
and glazed. Umbonal-ventral sulcus expressed internally as a weak to strong rib. Shell
with or without special laminae for the attachment of the posterior adductor muscle.

On the basis of shell characters the genus Jowannetia des Moulins is most closely re-
lated to Nettastomella Carpenter. The young shells of the two genera are almost insep-
arable, having a similar type of sculpture, umbonal reflection, pedal gape and umbonal-
ventral sulcus. Both lack the apophyses, and the type of siphonoplax is similar even
though restricted to a single valve in Jouannetia. The outstanding difference between
the two is the great production of the callum in Jowannetia.

The type of Jowannetia is a Tertiary fossil from the shell marls of Merignac near Bor-
deaux, France. The specimens, which were taken from coral, are unusually well pre-
served and show all the characteristics of the genus including the special laminae for the
attachment of the posterior adductor muscle. The most closely related living species,
Jouannetia cumingu Sowerby, from the Indo-Pacific, also bores into coral and soft coral-
line rock.

Sieverts (1933) reviewed the genus particularly as regards the fossil forms, recording
27 species, only 8 of which are recent. Of these eight, however, one is a synonym and
two are of doubtful value. She stated that the genus occurred as far back as the Upper
Cretaceous and was world-wide in distribution. Recent species, except for J. vignoni
Fischer from Gabon, West Africa, and J. qguillingi Turner from the Western Atlantic
are restricted to the Pacific.

Very little is known concerning the biology of this group. Fischer (1860, 1862) de-
scribed and figured the general morphology of J. cumingii Sowerby and J. globosa
Sowerby (=Jd. globulosa Quoy and Gaimard). These descriptions, based upon preserved
material, show the basic resemblance of this group to the other members of the Phola-
didae. A well illustrated account of the morphology of J. cumingu Sowerby by E. Egger
(1887) considers in detail the various systems of this species. However, though this spe-
cies is not excessively rare, as are most in this genus, nothing is known of its life history.
So far as is known all species are coral and rock-borers.

Genotype, Jouannetia semicaudata des Moulins, monotypic.
The genus Jouannetia is divided into two subgenera and these may be keyed as follows:

Shell with special laminae for the attachment of the posterior adductor
muscle. Margin of the siphonoplax smooth . . . . Jouannetia s.s.
Shell without special laminae for the attachment of the posterior adduc-
tor muscle. Margin of the siphonoplax pectinate . . . Pholadopsis

Subgenus Jouannetia des Moulins

Jouannetia des Moulins 1828, Bulletin d’Histoire Naturelle Société Linnéenne, Bordeaux 2, p. 244.

‘The species included in the subgenus Jowannetia are characterized by having special
laminae projecting inwardly from the posterior portion of the valves for the attachment

Eastern Pacific JOHNSONIA, No. 34 135

of the posterior adductor muscle, and in having the posterior margin of the siphonoplax
smooth.

There are only two living species in this subgenus: J. cumingii Sowerby from the Indo-
Pacific and J. duchassaingi Fischer from the Eastern Pacific.

Subgenotype, Jouannetia semicaudata des Moulins, monotypic.

Jouannetia (Jouannetia) duchassaingi Pischer
Plate 82

Jouannetia duchassaingi ‘Deshayes’ Fischer 1862, Journal de Conchyliologie 10, pp. 375-376, pl. 15, fig. 3
(Panama).

Distinctive characters. Shell reaching about 2 inches in length, having the callum longi-
tudinally striated, the posterior margin of the siphonoplax smooth, and having special
laminae for the attachment of the posterior adductor muscle.

Description. Shell white, globose, solid, reaching 50 mm. (about 2 inches) in length
and 41 mm. (about 14 inches) in height and having a callum in the adult stage. Young
shells nearly equivalved and widely gaping. Valves divided into two areas by an umbonal-
ventral sulcus which is very narrow on the right valve but wide on the left valve. Anterior
portion triangular in outline and sculptured by close-set concentric ridges and radial ribs
which are most clearly marked near the dorsal margin. Posterior area sculptured with fine
concentric ridges. Umbos prominent, located near the center of the dorsal margin. Pedal
gape closed in the adult by a thick, striated callum which extends dorsally over the beaks
and covers the anterior adductor muscle. Siphonoplax produced on the right valve only.
It is broad at the base and tapers to a narrowly rounded smooth extremity. Interior of
the valve smooth, umbonal-ventral sulcus expressed internally as a rib. Just posterior to
the sulcus there is a large lamina or shelf which forms the attachment area for the poste-
rior adductor muscle. A large shield over the umbos forms the attachment area of the
anterior adductor muscle.

The soft parts of this species are unknown.

length height ratio h:]
50 mm. 4] mm. 12 Holotype

Plate 82. Jouannetia duchassaingi Fischer. Figs. 1-2. Holotype (after Fischer, Journal de Conchyliologie
10, pl. 15, figs. $a—b). Fig. 1. Internal view of right valve showing the umbonal reflection and the large
shelf for the attachment of the posterior adductor muscle. Fig. 2. External view of the right valve showing
the smooth, triangular siphonoplax, the sculptured callum, and the extension of the callum over the dorsal
surface of the shell.

136 JOHNSONIA, No. 34 Jouannetia

Types. The type of Jouannetia duchassaingi Fischer was in the collection of Deshayes
and, according to Sherborn, is now probably in the British Museum. The type locality is
Panama (west coast).

Remarks. Until recently this species was known only from the original specimen and
description. Dr. A. A. Olsson, however, obtained it on a recent trip to Ecuador. His
specimens from Manta were beach worn, but probably had been boring in the shelving
ledges of shale which extend to the west of the city. This species is most closely related
to J. cumingu Sowerby but differs from it by being much larger and having the callum
longitudinally striated.

Range. From Panama south to Manta, Ecuador.

Specimens examined. Ecuapor: Manta (A. A. Olsson).

Subgenus Pholadopsis Conrad

Pholadopsis Conrad, June 1849, Proceedings Academy Natural Sciences Philadelphia 4, p. 156 (genotype,
Pholadopsis pectinata Conrad, monotypic); Conrad 1850, Journal Academy Natural Sciences Philadelphia (2)
1, p. 279: Tryon 1862, Proceedings Academy Natural Sciences Philadelphia, p. 216.

Triomphalia Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, p. 500 (genotype, Triomphalia globosa Sow-
erby | =P. globulosa Quoy and Gaimard |, subsequent designation, Stoliczka 1870); Sowerby 1849 [1850] Pro-
ceedings Zoological Society London, p. 160.

Triomphala ‘Sowerby’ Clessin 1892, Conchylien-Cabinet 11, part 4, p. 35 [error for Triomphalia Sowerby ].

The species included in the subgenus Pholadopsis Conrad are characterized by having
the siphonoplax pectinate and by having the posterior adductor muscle inserted in the
normal position. In addition, there is a small mesoplax which in the adult form is fused
to the callum.

This subgenus has been referred to by numerous authors as T7riomphalia Sowerby:
however, as a footnote to his original description, Sowerby mentions the genus Phola-
dopsis and states that it may be identical with 77iomphalia. Consequently, though both
Conrad’s and Sowerby’s work appeared in 1849, there seems little doubt as to which
name has priority. Lamy (1926) curiously uses 7’riomphalia rather than Pholadopsis, but
when considering the species included in the subgenus, he places pulcherrima Sowerby
in the synonymy of pectinata Conrad. These two species were described by Sowerby and
Conrad at the same time as their respective genera. Recent workers following Lamy have
given priority to Sowerby for the genus and to Conrad for the species.

All four species in this subgenus are exceedingly rare. Each is very distinctive and
easily identified. There is one species in each of the four major oceanic divisions; J. glo-
bosa Sowerby in the Indo-Pacific; J. pectinata Conrad in the Eastern Pacific: J. quil-
lingi Turner in the Western Atlantic, and J. vignoni Fischer in the Eastern Atlantic.
This last-named species from Gabon, West Africa is known only from the original de-
scription and figures (Fischer 1862, Journal de Conchyliologie 10, p. 276, pl. 15, fig. 4).
It has not been found since and the location of the types is unknown. However, the
figures are excellent and there is no question as to the uniqueness of this species.

Subgenotype, Pholadopsis pectinata Conrad, monotypic.

Eastern Pacific JOHNSONIA, No. 34 137

Jouannetia (Pholadopsis) pectinata Conrad
Plate 83

Pholadopsis pectinata Conrad, June 1849, Proceedings Academy Natural Sciences Philadelphia 4, p. 156
(Baja California and Peru); Conrad 1850, Journal Academy Natural Sciences Philadelphia (2) 1, p. 279, pl.
39, fig. 3.

Triomphalia pulcherrima Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, p. 501, pl. 106, figs. 58-59 (in
soft rock at low water, West Colombia); Sowerby 1849 [1850], Proceedings Zoological Society London for
1849, p. 161, pl. 5, figs. 2a—d.

Jouannetia pectinata Conrad, Gray 1851, Annals Magazine Natural History (2) 8, p. 383.

Jouannetia (Pholadopsis) pectinata Conrad, Tryon 1862, Proceedings Academy Natural Sciences Philadelphia
15, p. 216.

Jouannetia ( Triomphalia) pectinata Conrad, Lamy 1926, Journal de Conchyliologie 69, p. 219; Sieverts 1933,
Neues Jahrbuch fiir Mineralogie, Geologie und Palaeontologie 71, Beilage-Band Abt. B, pt. 2, p. 289; Hert-
lein and Strong 1950, Zoologica 35, no. 19, p. 248.

Distinctive characters. Shell with a greatly produced and irregularly sculptured callum.
Posterior slope sculptured with smooth concentric ridges and growth lines. Posterior
margin of the siphonoplax finely pectinate.

Description. Adult shell pear-shaped in outline, inequivalve, reaching about 51 mm.
(about 2 inches) in length and 383 mm. (about 14 inches) in height, white in color, thin,
fragile, and with a greatly produced callum. Young shell equivalve, beaked and widely
gaping anteriorly; closed posteriorly. Shell divided into two distinct parts by a narrow
umbonal-ventral sulcus. Anterior portion triangular in outline and sculptured by numer-
ous imbricated, concentric ridges and radial ribs. There is a marked constriction of the
shell at the umbonal-ventral sulcus. Sulcus narrow and sculptured with concentric ridges
and growth lines. Posterior margin of the sulcus indefinite, especially on the right valve.
Posterior slope sculptured with concentric ridges and growth lines. Umbos prominent
and located near the center of the dorsal margin. Umbonal reflection narrow, free and
high. Pedal gape closed in the adult shell by a callum which is nearly typical on the
right valve but greatly enlarged on the left valve so that it overlaps the right valve. The
callum extends dorsally between the beaks and the left valve enlarges to form the covering
of the anterior adductor muscle. Ventrally the callum of the right valve stops at the

Plate 83. Jouannetia pectinata Conrad. Fig. 1. Dorsal view of the paratype showing the greatly enlarged,
overlapping callum, the small mesoplax which has been displaced by the growth of the callum, and the
pectinate siphonoplax of the right valve. Fig. 2. Internal view of the left valve of the holotype showing
the tremendous callum, the small posterior adductor muscle scar, the small chondrophore and the lack of
the apophysis. Fig. 3. External view of the right valve of the holotype showing the much smaller callum
of this valve and the pectinate siphonoplax (all 14x).

138 JOHNSONIA, No. 34 Jouannetia

umbonal-ventral sulcus but on the left valve additional material is added along the ven-
tral and posterior margins of the posterior slope. Right valve elongated posteriorly, form-
ing the siphonoplax which is broad at the base and tapers to a narrowly rounded, finely
pectinate posterior margin. Mesoplax small, triangular and sculptured with concentric
growth lines. In the adult shell it is fused to the callum, and the enlarging of the cal-
lum displaces it so that it is set at an angle to the longitudinal axis of the shell.

Interior of the shell white and glazed. Umbonal-ventral sulcus expressed internally as
a ridge. Anterior adductor muscle scar covering the umbonal reflection. Posterior ad-
ductor muscle scar small, elongate-oval, and located high on the posterior slope. Pallial
sinus profound, extending anterior to the umbonal-ventral ridge. Chondrophore on the
left valve small. The soft parts of this species are unknown.

length * height ratio h:]
51.5 mm. 33.5 mm. Lis Holotype
41.5 28.5 1.4 Paratype
28.0 Dike 1.3 Port Parker, Costa Rica

* The measurements are for entire adult specimens including the callum and the siphonoplax.

Types. The holotype of Pholadopsis pectinata Conrad is in the Academy of Natural
Sciences Philadelphia, no. 51075. The type locality as restricted by Hertlein and Strong
(1950) is the east coast of Baja California. The type of Triomphalia pulcherrima Sow-
erby is in the British Museum; the type locality is West Columbia.

Remarks. Jouannetia pectinata Conrad is a very distinctive species readily distinguished
from all others in this subgenus by its large size, its fragile shell and its greatly enlarged,
irregularly sculptured callum. It is most closely related to J. quillingi Turner of the
Western Atlantic. See Remarks under quillingi. It is also closely related to J. globulosa
Quoy and Gaimard from the Indo-Pacific. This latter is a much smaller species with a
broad umbonal-ventral sulcus and a globose rather than pear-shaped outline in the adult
stage. From J. vignoni Fischer of West Africa it differs by its much larger size and pro-
portionately much smaller siphonoplax. In addition, the siphonoplax of J. vignoni has a
few broad, coarse pectinations while those of J. pectinata are numerous and small. Speci-
mens of J. pectinata reach 50 mm. in length while those of J. globulosa average 23 mm.
and the type of J. vignoni is 10 mm. in length. However, in this group as well as all
members of the Pholadidae, size alone should not be depended upon as a criterion for
separating species.

Nothing is known of the biology of this rare species. The specimens collected by Dr.
Thomas B. Wilson and sent to the Academy of Natural Sciences, Philadelphia were alive
when collected and were probably taken in the intertidal zone. The specimens collected
by Hugh Cuming were in soft stone at low water.

Range. From Cedros Island, Baja California south to Peru (Conrad, 1849).

Specimens examined. Mexico: Baja California (ANSP). Eight miles west of Cedros
Island (N. Lat. 28°05’: W. Long. 115°31’) in 64 fathoms (Hancock Foundation). Cos'ra
Rica: Port Parker (California Acad. Sci.). Ecuapor: South of Punta Santa Elena
(S. Lat. 2°12’; W. Long. 81°00’) (Hancock Foundation).

Western Atlantic JOHNSONIA, No. 34 139

Jouannetia (Pholadopsis) quillingi, new species
Plate 84

Distinctive characters. Shell with a large and irregularly sculptured callum. Posterior
slope with high, thin, concentric ridges bearing recurved spines. Siphonoplax small and
with few, spine-like pectinations.

Description. Adult shell nearly globular in outline, inequivalve, reaching about 21
mm. (about £ of an inch) in length and 15 mm. (about 2 of an inch) in height, white in
color, thin, fragile and with a greatly produced callum. Young shells equivalve, beaked
and widely gaping anteriorly and closed posteriorly. Valves divided into two distinct
parts by the umbonal-ventral sulcus. Anterior portion triangular in outline and sculp-
tured with numerous laminated, imbricate, concentric ridges and weak radial ribs which
are indicated only by the radial arrangement of the imbrications. There is a marked
constriction of the shell at the umbonal-ventral sulcus. ‘The sulcus itself is very narrow
and appears only as the line of constriction of the shell. Posterior slope sculptured with
thin, raised, concentric ridges and growth lines. The ridges are scalloped and on perfect
specimens bear long recurved spines. Umbos prominent and located near the anterior
third of the specimen. Umbonal reflections narrow, free and raised well above the surface
of the shell. Pedal gape closed in the adult shell by a callum which is nearly typical on
the right valve but is enlarged on the left valve so that it overlaps the right valve. The
callum extends dorsally between the beaks and on the left valve is enlarged to form the
covering for the anterior adductor muscle. Ventrally the callum of the right valve stops
at the sulcus, while on the left valve, additional material is added along the ventral and
posterior margin of the posterior slope. Right valve slightly elongate posteriorly, form-
ing the small siphonoplax which is strongly pectinate on its posterior margin. There are
fifteen strong, spine-like pectinations on the holotype. Mesoplax small, wedge-shaped
and sculptured with strong, chevron-shaped growth lines. The enlarging of the callum
displaces the mesoplax in the adult so that it is set at an angle to the longitudinal axis
of the shell.

Interior of the shell white and glazed. Umbonal-ventral sulcus expressed internally as
a narrow ridge. Muscle scars barely visible on the specimens examined. Posterior adduc-

Plate 84. Jouannetia quillingi Turner. Holotype from Lake Worth, Florida. Fig. 1. External view of the right
valve showing the pectinations on the siphonoplax and the pronounced sculpture of the posterior slope.
Fig. 2. Internal view of the left valve showing the small chondrophore and the smooth posterior margin.
The dorsal portion of the callum is broken, but in complete specimens it is similar to that shown in plate
83, fig. 2 (about 23x).

140 JOHNSONIA, No. 34 Jouannetia

tor muscle scar small, oval in outline and located high on the posterior slope. Anterior
adductor muscle scar covering most of the umbonal reflection. Pallial sinus extending
anteriorly to the umbonal-ventral ridge. Ligament small and internal. There is a small
chondrophore on the left valve. Soft parts of this species are unknown.

length * height ratio h:1]
21.5 mm. 15e5 mm, 1.38 Holotype
16.5 14.0 1.18 Pawleys Island, South Carolina
9.0 5.5 1.60 off Cape Fear, North Carolina

* The measurements are for entire adult specimens including the callum and the siphonoplax.

Types. The holotype from the north end of Lake Worth Inlet, Lake Worth, Florida
is in the Museum of Comparative Zoédlogy, no. 1899138. Paratypes are in the collection
of Tomand Paul McGinty, Boynton Beach, Florida. The types were collected by Paul
McGinty and Ben Quilling. We take pleasure in naming this species for Mr. Quilling.

Remarks. This is the first record for the genus Jouannetia in the Western Atlantic.
This new species is close in its relationship to J. pectinata Conrad of the Eastern Pacific
but may be readily distinguished from it by its much smaller size, by the pronounced,
spiny sculpture on its posterior portion and by the relatively few large pectinations on
the siphonoplax. From J. vignoni Fischer of West Africa it is differentiated by its short,
broad siphonoplax which makes the right valve only slightly longer than the left in adult
specimens. In J. vignoni the siphonoplax is long and narrow, the right valve of the adult
specimen being nearly twice the length of the left valve. In addition, the pectinations
on the siphonoplax of vignoni are broad and coarse while those of quillingi are long and
spine-like.

Little is known of the ecology of this species. The McGintys have written that they
found it “‘living in a large piece of submerged and rotted wood.”’ In 1885 the Albatross
obtained specimens while dredging in 18 fathoms off Cape Fear, North Carolina; how-
ever, no record was made as to whether the specimens were living or dead, or in what
they were found. At Pawleys Island, South Carolina, this species was apparently living
in the intertidal zone.

Woodring (1925, Miocene Mollusks from Bowden, Jamaica, p. 198, pl. 26, figs. 18-
15) describes a Jouannetia sphaeroidalis Guppy MS. In his remarks Woodring states
that if this is a Jowannetia the specimens are young because there is *‘. . . no trace of a
posterior appendage on the right valve.’’ At the same time he describes the anterior callus
or callum which would indicate that they were not young. At the time the callum is
produced, all other adult features are also added, a condition which exists throughout all
of the Pholadidae. Therefore, from the description and figures of Woodring, it seems
clear that this is not a Jowannetia. It is perhaps a Diplothyra.

Range. From off Cape Fear, North Carolina south to Lake Worth, Florida.

Specimens examined. Norra Carouina: Albatross, station 2615, 27 miles east southeast
of Cape Fear (N. Lat. 33°45’; W. Long. 77°25’) in 18 fathoms; Albatross, station 2161,
25 miles east southeast of Cape Fear (N. Lat. 38°42’; W. Long. 77°31’) in 17 fathoms
(both USNM). Sourn Carotina: Pawleys Island (Charleston Museum). FLoripa:
North end of Lake Worth Inlet, Lake Worth, Florida (T. McGinty); off Naples in
4 to 6 fathoms (‘T. L. Moise).

Eastern Pacific JOHNSONIA, No. 34 141

Genus Nettastomella Carpenter

Netastoma Carpenter 1864, Report British Association Advancement of Science for 1863 [1864], p. 637
(genotype, Netastoma darwinii ‘Sowerby’ Carpenter); non Nettasloma Rafinesque 1810.

Nettastomella Carpenter 1865, Proceedings Zoological Society London, p. 202 [new name for Netastoma
Carpenter; non Neftastoma Rafinesque|; Hertlein and Strong 1950, Zoologica 35, p. 247.

Netastomella ‘Carpenter’ Paetel 1875, Familien-und Gattungsnamen der Mollusken, Berlin, p. 138 [error
for Nettastomella Carpenter].

Nettastoma ‘Carpenter’ Lamy 1926, Journal de Conchyliologie 69, p. 155 [error for Nelastoma Carpenter].

Shell small to moderate in size, gaping widely anteriorly, the pedal gape not closed in
the adult stage by a calcareous callum. In adult specimens a narrow band of calcareous
material is laid down anterior to the imbricated area and this may be fluted or simply
marked by growth lines. A chitinous callum covers the remainder of the pedal gape with
only a minute central pore remaining open. The foot atrophies in the adult. Umbonal
reflection sculptured or smooth, simple, narrow and free for most of its length. Dorsal
plates lacking, siphonoplax short and widely diverging or long and acuminate. Apophy-
ses lacking.

At the time Carpenter described his genus Netastoma he was working with the fauna
of the west coast of North America and incorrectly determined young specimens of N.
rostrata Valenciennes as N. darwinit Sowerby. This was an easy error to make as young
specimens of N. rostrata, when they first begin to produce the siphonoplax and before
they have begun to elongate it, look much like specimens of N. darzwinii as illustrated by
Sowerby, though they are much smaller. However, adult specimens of rostrata are not
only smaller in size but eventually produce a long, acuminate siphonoplax, while the
South American species produces short, greatly thickened and widely diverging out-
growths on the posterior margin of the shell.

Through the kindness of Dr. Katherine VanWinkle Palmer we have been able to ex-
amine specimens from the Carpenter Collection now in the Redpath Museum, McGill
University, Montreal, Canada. In this collection there are four lots of Nettastomella ros-
trata Valenciennes all labeled in Carpenter’s own hand as Nettastomella darwini Sowerby.
Consequently, there seems to be little doubt as to Carpenter’s confusion regarding these
two species. In his brief description of the genus, Carpenter refers to the strong ‘‘frills””
on the posterior portion of the shell and to the valves being ‘“‘prolonged, like duck’s
bill.’’ This applies exactly to rostrata, the West Coast species, while on darwini Sowerby
the posterior slope is marked only by low, indistinct ridges and growth lines and the si-
phonoplax is short and diverging. It seems logical, therefore, even though darwini
remains in this genus, that the type of Nettastomella becomes N. darwini ‘Sowerby’
Carpenter (=rostrata Valenciennes); non darwini Sowerby.

Only two species are known in this genus: N. rostrata Valenciennes, occurring along
the California coast and N. darwinitt Sowerby which is restricted to Chile, Argentina
and Uruguay.

Genotype, Netastoma darwinit ‘Sowerby’ Carpenter 1864 (= N. rostrata Valenciennes),
monotypic; non Pholas darwinii Sowerby 1849).

142 JOHNSONIA, No. 34 Nettastomella

Key to the species of Nettastomella

1. Shell small, never over an inch in length (not including the
siphonoplax), siphonoplax in adult specimens long and thin,
not diversing 0-3. s + 40. set 2 eee eee

ins)

Shell large, reaching 14 inches in length, siphonoplax heavy,
short and widely diverging . . . .... . . N.darwinii

Nettastomella darwinii Sowerby
Plates 85-86

Pholas darwinii Sowerby 1849, Thesaurus Conchyliorum 2, part 10, p. 490, pl. 107, figs. 76-77 (Chiloe
(Island, Chile], C. Darwin, collector); Sowerby 1873, Conchologica Iconica 18, pl. 4, fig. 12. [Not described
in the Proc. Zool. Soc. London 1849 as indicated in the Thesaurus. |

Pholadidea (Nettastomella) darwinii Sowerby, Oldroyd 1924, Stanford University Publications, University
Series, Geological Sciences 1, p. 213, pl. 42, figs. 7-8; A. Carcelles 1950, Anales del Museo Nahuel Huapi
2, p. 82, pl: 5; fig..93.

Distinctive characters. Shell 1 to 14 inches in length, very widely gaping anteriorly,
closed posteriorly and having a short, heavy, widely diverging siphonoplax. Callum ex-
isting as a narrow band which is devoid of sculpture. Sculpture on the posterior slope
consisting of low concentric ridges and growth lines.

Description. Shell reaching 37 mm. (about 14 inches) in length and 21 mm. (about £
of an inch) in height, white, rather light in structure, beaked anteriorly and widely gap-
ing; closed posteriorly. Adult with a partial callum and a short, heavy, widely diverging
siphonoplax. Valves divided into two areas by a marked umbonal-ventral constriction
though no true sulcus is formed. Anterior portion triangular in outline and sculptured
with thin, high, concentric ridges and weak radial ribs. Posterior portion sculptured only
with low ridges and growth lines. Umbos prominent and located near the anterior third
of the shell. Umbonal reflection narrow and free. Callum in the adult specimen existing
as a narrow band of unsculptured calcareous material extending along the anterior mar-
gin and dorsally over the beaks to the umbo. Siphonoplax short, calcareous, solid in
structure and widely diverging. So far as known there are no other accessory plates.

Plate 85. Nettastomella darwinii Sowerby. Figs. 1-3. Holotype of Pholas darwinii Sowerby (= Nettastomella
darwiniti Sowerby). Chiloe Island, Chile. Fig. 1. Dorsal view showing the umbonal reflection and the dor-
sal extension of the callum. Fig. 2. Ventral view showing the chondrophore. Fig. 3. Side view showing
the narrow band of the calcareous portion of the callum (all 2x).

Photographs received through the kindness of Mr. G. L. Wilkins, British Museum (Natural History).

Western Atlantic JOHNSONIA, No. 34 143

Interior of shell white and glazed. Muscle scars well marked; pallial sinus not evident in
the few specimens examined. Apophyses lacking. Chondrophore small. Preserved ma-
terial being unavailable, it is impossible to state whether or not the remainder of the pedal
gape is closed by a chitinous callum.

length height ratio h:]

37.0mm. 21.0mm. hey) Piriapolis, Maldonado, Uruguay

85.0 21.0 1.6 a a ip

35.0 922.0 1.5 Comodoro, Rivadavia, Chubut, Argentina

Types. The holotype of Pholas darwini Sowerby is in the British Museum (Natural
History). The type locality is Chiloe Island, Chile, C. Darwin collector.

Remarks. Nettastomella darwini Sowerby is apparently a very rare species and has
been confused with N. rostrata Valenciennes from the coast of California, probably be-
cause of the early use of this name for the Californian species by P. P. Carpenter. How-
ever, N. darwinit can readily be told from N. rostrata by its larger size, its relatively
smooth posterior portion and callum, and its short, heavy, widely diverging siphonoplax.

Range. From Maldonado, Uruguay south to Chubut, Argentina and north along the
coast of Chile to Chiloe Island.

Records. Urucuay: Piriapolis, Maldonado (A. Carcelles). ARGENTINA: Zona Mili-
tar de Comodoro Rivadavia (A. Carcelles); Mar del Plata(USNM). Cuine: Canal de
Chacao, north end of Chiloe Island in 22 fathoms (Univ. of Lund Exped. ).

Plate 86. Nettastomella darwinii Sowerby. Figs. 1, 3 and 4. Zona Militar de Comodoro Rivadavia, Argentina.
Fig. 2. Piriapolis, Maldonado, Uruguay (all about 14x).

Nettastomella rostrata Valenciennes
Plate 87

Pholas rostrata Valenciennes 1846 [in] Abel du Petit-Thouars, Voyage Autour du Monde sur la Frégate La
Vénus. Atlas de Zoologie, Mollusques, pl. 24, fig. 4 [name listed on plate; no locality given]; Lamy 1921,
Bulletin Muséum National d’ Histoire Naturelle, Paris 27, p. 182 (Monterey, California).

Nettastomella darwinii ‘Sowerby’ Carpenter 1865, Proceedings Zoological Society London, p. 202 (Monterey

and San Diego [California]; Vancouver [ British Columbia | ).

144 JOHNSONIA, No. 34 Nettastomella

Pholadidea ( Nettastomella) rostrata Valenciennes, Oldroyd 1924, Stanford University Publications, Univer-
sity Series, Geological Sciences 1, no. 1, p. 213, pl. 13, figs. 10-11, pl. 38, figs. 7, 10.
Nettastomella rostrata Valenciennes, Dall 1921, Bulletin 112, United States National Museum, p. 56.

Distinctive characters. Shell small, # inch or less in length, widely gaping anteriorly,
closed posteriorly and with a long, tapering siphonoplax in the adult stage. Callum ex-
isting as a band which is sculptured with high, thin flutes. Sculpture on the posterior
slope consisting of thin, high, concentric ridges.

Description. Shell small, reaching about 20.1 mm. (about } inch) in length and 7.8
mm. (about 4 inch) in height, fragile, beaked anteriorly and widely gaping in the young
stage; closed posteriorly. In the adult stage there is a long, acuminate siphonoplax. Shell
divided into two areas by a marked constriction of the valves, though no true sulcus is
formed. Anterior portion triangular in outline and sculptured with numerous, thin, high,
concentric ridges and weak radial ribs. Rounded imbrications are formed where the radial
ribs cross the ridges. Posterior portion sculptured with high, thin, concentric flutes.
Umbos prominent, located near the anterior third of the shell. Umbonal reflection nar-
row and free. The calcareous portion of the callum exists as a narrow band extending
along the anterior margin of the beaks and dorsally over the umbonal reflections to en-
close partially the anterior adductor muscle. The callum is very fragile and is sculptured
with high, thin flutes. The remainder of the pedal gape is closed by a chitinous covering
leaving only a minute central pore open in the fully adult specimen. Siphonoplax vary-
ing in length and width depending upon the age of the specimen and the depth to which
it is boring. In young specimens it is very short and broad, and may be reflected out-
wardly, but as the specimen grows older, the siphonoplax elongates, becoming acuminate
and often rather irregular in shape. There are no other accessory plates. Interior of the
shell white and glazed. Posterior adductor muscle scar large, oval in outline and set high
on the posterior slope. Pallial sinus extending anterior to the umbonal-ventral ridge.
Apophyses lacking. Siphons slightly papillose at the posterior end, white in color and
with a periostracal sheath only at the base. ‘They may be extended a short distance be-
yond the end of the siphonoplax or contracted completely within the shell. Foot large,
round and truncated in young specimens; atrophied in the adult.

Plate 87. Nettastomella rostrata Valenciennes. Fig. 1. San Diego, California. Dorsal view showing the exten-
sion of the marginal callum over the beaks, the strong sculpture on the posterior slope and the siphonoplax.
Fig. 2. San Pedro, California. A small but adult specimen showing the very small anterior portion, the
wide band of the callum and the greatly extended siphonoplax. Figs. 3-4. Laguna, California. Fig. 3.
Ventral view of a young specimen just beginning to produce the siphonoplax. Fig. 4. Internal view show-
ing the cavity formed by the reflection of the callum over the beaks, and the deep pallial sinus. Fig. 5. San
Pedro, California. Side view showing the sculpturing on the callum and the free umbonal reflection (all 23x),

Eastern Paciftic JOHNSONIA, No. 34 145

length height ratio h:]

=)

12.5mm.* 12.0mm. ile White Point, San Pedro, California

9.8 8.1 2 Laguna, California
OTD 7.9 ae San Pedro, California
8.2 5.5 1.4 = . =

* Not including the siphonoplax

Types. According to Lamy 1921, Bulletin Muséum National d’ Histoire Naturelle,
Paris 27, p. 182, the type of P. rostrata Valenciennes is in the Paris Museum. The type
locality, Monterey, California, though not published by Valenciennes, is on the original

label.

Remarks. Nettastomella rostrata Valenciennes is one of the smallest of the rock-boring
pholads. It is found in soft shale beds along the California coast. Specimens sent by E. P.
Chace were from soft mud-shale blocks brought in near shore at White Point, San Pedro,
California; others have been dredged in depths up to 55 fathoms. They are not deep
borers and their burrows are seldom perfectly straight. The siphonoplax, following the
contours of the burrow, may be very irregular in shape and curvature.

Nettastomella rostrata Valenciennes is not closely related to any other species. N. dar-
win, with which it has been confused, is quite a different species.

Range. Puget Sound (Dall 1921) and Vancouver (P. P. Carpenter 1865) south to off
Cedros Island, Baja California.

Specimens examined. CALIFORNIA: Duxbury Reef, Bolinas (USNM): Monterey
(MCZ; Redpath Museum; USNM): off Del Monte, Monterey Bay in 12 fathoms
(S. S. Berry); Moss Landing, Monterey Bay in 16 fathoms (MCZ); Santa Barbara
(MCZ:; Redpath Museum); White Point, San Pedro (KE. P. Chace; J. E. Fitch); San
Pedro (EK. P. Chace; USNM); Long Beach (USNM); Laguna (MCZ); Portuguese
Bend (N. Lat. 33°44’; W. Long. 118°22’) (Hancock Foundation); La Jolla (USNM);
San Diego (MCZ: Redpath Museum; USNM). Mexico: 15 miles off north end of
Cedros Island, Baja California (N. Lat. 28°23’: W. Long. 115°11’) in 55 fathoms (Han-
cock Foundation).

Subfamily Xylophaginae
Genus Xylophaga Turton

Xylophaga Turton 1822, Conchylia Insularum Britannicarum, p. 253; Purchon 1941, Journal Marine Bio-
logical Association United Kingdom 25, pp. 1-39.

Xylotomea Dall 1898, Transactions Wagner Free Institute of Science, Philadelphia 3, pt. 4, p. 821 (new
name for Xylophaga Turton non Xyloghagus Meuschen 1778). [This name is unnecessary however, as Xylophaga
and Xylophagus are not considered homonyms. Meuschen’s work has now been rejected by the International
Commission—Opinion 260].

Xilotoma ‘Dall’ Gigoux 1934, Revista Chilena de Historia Natural, Santiago 38, p. 285 (error for Xy/oto-
mea Dall).

Protoxylophaga Taki and Habe 1945, Venus 14, p. 112 (genotype, NXylophaga tomlini Prashad, original
designation).

Neoxylophaga Taki and Habe 1950, Illustrated Catalogue of Japanese Shells, no. 7, p. 46 (genotype, Xy/o-
phaga rikuzenica Taki and Habe, original designation).

Metaxrylophaga Taki and Habe 1950, Illustrated Catalogue of Japanese Shells, no. 7, p. 47 (genotype,
Metaxylophaga supplicata Taki and Habe, monotypic).

146 JOHNSONIA, No. 34 Xylophaga

Shell small, globose, teredo-like, beaked and widely gaping anteriorly throughout life,
broadly rounded and closed posteriorly. Beaks truncated at nearly right angles to the
anterior margin of the disc. Beaks and anterior portion of the dise sculptured with nu-
merous, finely denticulated ridges. Umbonal-ventral sulcus present but variable and ex-
pressed internally as a pronounced ridge which usually terminates in a small ventral con-
dyle. Umbonal reflection narrow and variable; it may or may not extend over the umbos.
Apophyses absent. Chondrophore and internal ligament present. Dorsal plates consisting
of a divided mesoplax which may cover the umbos or be placed anterior to them. Siphons
variable; in some species the excurrent siphon is truncated at the posterior margin of the
valves, in others it is nearly as long as the incurrent siphon.

Purchon (1941) removed the genus Yylophaga from the family Pholadidae and created
for it a new family, the Nylophaginidae. However, so little is known concerning the
soft anatomy of the many species of the Pholadidae, it seems unwise to make such a move
at this time. Certainly, on the basis of shell characters, Yylophaga is more closely related
to the Pholadidae than to the Teredinidae. The presence of the mesoplax, the lack of
apophyses, and the agglutinization of the fecal pellets to form a ‘‘chimney”’ are characters
found in other pholad genera. Many species of pholads have a small chondrophore and
internal ligament, and two genera, Nettastomella and Jouannetia, lack apophyses. Pur-
chon states that Yy/ophaga is a wood borer while the pholads are rock borers. However,
Martesia, one of the most abundant of the pholads, is also a wood borer. The alternation
of sexes which is mentioned as showing an affinity with the Teredinidae has also been
demonstrated for Martesia (Moore 1947). Though at least some species of Yylophaga
retain their young, this is of no great importance for in the Teredinidae some groups, such
as Teredo, do, while others, such as Bankia, do not. The reduction of the excurrent siphon
is mentioned as one of the characteristics peculiar to the Nylophaginidae, but in Xy/lo-
phaga atlantica Richards, the excurrent siphon is only very slightly shorter than the in-
current siphon and is in other respects normal. Consequently, we are including X ylophaga
in the Pholadidae in the subfamily Nylophaginae.

Three recently described genera have been submerged under the genus Xylophaga in
the synonymy given above. Fortunately it has been possible to study type specimens of
the type species of all three.

The genus Xylophaga seems to be largely pelagic, occurring mainly in floating and
waterlogged wood. The genus is world-wide in distribution, but does not occur in suffi-
cient numbers to be of any real economic importance. One species, X. abyssorum Dall,
has been recorded as boring into mud and soft stone, but this is open to question.

Genotype, Teredo dorsalis Turton, monotypic.

Xylophaga dorsalis Turton
Plate 88

Teredo dorsalis Turton 1819, A Conchological Dictionary of the British Islands, p. 185 (Devonshire Coast,
England).

Xylophaga dorsalis Turton 1822, Conchylia Insularum Britannicarum, p. 253; Verkrizen 1872, Norwegen
Seine Fjorde Und Naturwunder, p. 35; Purchon 1941, Journal Marine Biological Association United King-
dom 25, pp. 1-39.

Pholas xilophaga Deshayes 1835 [in] Lamarck, Animaux sans Vertébres, ed. 2, 6, p. 47.

Eastern Atlantic JOHNSONIA, No. 34 147

Distinctive characters. Beak extending less than half the distance to the ventral mar-
gin. Mesoplax large, divided, semicircular in outline and placed over the umbos. Pos-
terior adductor muscle scar more or less circular in outline and nearly smooth. Umbonal-
ventral sulcus shallow. Excurrent siphon truncated at the posterior margin of the valves.

Description. Shell globose, small, reaching about 10 mm. (about 2 inch) in length,
white to light brown in color and fragile. Shell gaping anteriorly throughout life, closed
posteriorly. Anterior slope beaked, the beak extending less than half the distance to the
ventral margin and truncated at nearly a right angle, giving the shell a teredo-like ap-
pearance. Beak sculptured with numerous rows of finely denticulated ridges which run
parallel to the ventral margin. Anterior portion of the disc sculptured with close-set den-
ticulated ridges which parallel the anterior margin. The ridges of the disc are much more
closely set than on the beak. Posterior to the denticulated portion of the disc there is
a broad, shallow umbonal-ventral sulcus which is sculptured only with fine growth lines.
Posterior slope inflated, with a high, ear-like lobe posteriorly and sculptured only with
growth lines. Umbos prominent and located near the anterior fourth of the shell. Um-
bonal reflections narrow, only partially covering the umbos, appressed near the umbos
and free anteriorly. Dorsal plates consisting of a relatively large, divided, semicircular
mesoplax which covers the umbos. Interior of shell white and glazed. Muscle scars well
marked. Posterior adductor muscle sear large, smooth and nearly circular. Anterior ad-
ductor muscle scar covering most of the umbonal reflection. Pallial sinus not apparent.
Umbonal-ventral sulcus expressed internally as a pronounced ridge with a condyle at the
ventral margin. Chondrophore present in left valve. Internal ligament present but small.
Apophyses lacking.

Siphons small, the incurrent siphon extending about twice the length of the shell, and
as much as 1 cm. beyond the surface of the wood. Aperture of the incurrent siphon sur-

Internal view of valve to show the chondrophore, the pronounced

Plate 88. Xylophaga dorsalis Turton. Fig. 1.
2, External view of valve showing the

umbonal-ventral ridge and the posterior adductor muscle sear. Fig.
shallow umbonal-ventral suleus. Fig. 3. Anterior view (pedal gape) of apposed valves to show the chon-
drophore and the points of contact of the valves. Fig. 4. Dorsal view to show the mesoplax in place. Fig.
5. Ventral view of the mesoplax. Fig. 6. Dorsal view of the mesoplax. All from Millport, Ayr, Scotland.

148 JOHNSONIA, No. 34 Xylophaga

rounded by six small cirri. Excurrent siphon truncated at about the posterior margin of
the valves, and lacking cirri. A paired series of lappets, all that remains of the excurrent
siphon, extend from the truncation of the excurrent siphon along the dorsal surface of
the incurrent siphon. Fecal pellets are accumulated in the posterior portion of the bur-
row and these are fused together to form a ‘‘chimney”’ similar to that found in Parapholas.

length height ratio h:]
6.5 mm. Heap ionic i}, Plymouth, England
5.0 5.0 1.0 v is

Types. The types of Xylophaga dorsalis Turton are probably in the British Museum.
The type locality is Devonshire Coast, England.

Remarks. A description of this European species is included here; first, because it is
the type of the genus and secondly, because until 1942 our Western Atlantic species was
confused with it. The two species, however, are quite distinct and can be easily separated
by the shape and position of the mesoplax. In _Y. dorsalis Turton, this plate is large,
conspicuous, composed of two ear-shaped parts and covers the umbos. In _X. atlantica
Richards the mesoplax is small, composed of two triangular parts and is located anterior
to the umbos.

Xylophaga dorsalis is most closely related to_Y. globosa Sowerby from the west coast
of South America. It differs from this species mainly in the shape of the mesoplax and
in the more pronounced umbonal-ventral sulcus of g/obosa.

Xylophaga dorsalis is not a common species, but on occasion it may be abundant in
restricted localities. It occurs mainly in floating or submerged waterlogged wood, but
occasionally is found in water-front structures, buoy markers and submarine cables, caus-
ing a minor amount of damage. Like other members of this family, specimens of this
species boring into coverings of cables, hard wood or other similar substrata are malformed
and stenomorphie.

A fine paper by R. D. Purchon on the anatomy and biology of Yylophaga dorsalis
appeared in 1941. In this paper, Purchon considers in detail the ciliary mechanism of the
mantle cavity, the morphology of the digestive and reproductive systems, and general
shell structure. He demonstrated sexual dimorphism in this species, described the vesic-
ula seminalis for the storage of sperm, and suggested that self-fertilization would be of
great survival value to a pelagic species which lives in isolated communities.

Range. From Lofoten Islands, Norway south to the western Mediterranean, the
Canary and Madeira Islands.

Specimens examined. Norway: below Drébak, Oslofjord (USNM). Scorianp: off
Shetland Islands (USNM); Millport, Ayr (C. M. Yonge). ENGLanp: Plymouth
(USNM). France: La Croisic, Loire-Inférieure (MCZ).

Xylophaga globosa Sowerby
Plate 89

Xylophaga globosa Sowerby 1835, Proceedings Zoological Society London, p. 110 (Valparaiso, Chile, from
piece of wood dredged at 100 fathoms); Sowerby 1849, Thesaurus Conchyliorum 2, pt. 10, p. 503, pl. 108,
figs. 101-102.

Pholas gibbosa d’Orbigny 1846, Voyage dans L’ Amérique Méridionale 5, pt. 3, p. 501.

Xylotomea globosa Sowerby, Dall 1909, Proceedings United States National Museum 37, p. 277.

Xilotoma globosa Sowerby, Gigoux 1934, Revista Chilena de Historia Natural, Santiago, p. 285.

Eastern Pacific JOHNSONIA, No. 34 149

Distinctive characters. Beak extending at least half the distance to the ventral margin.
Mesoplax large, divided and covering the umbos. Posterior adductor muscle scar elongate-
oval in outline and with a shelf at its anterior margin. Umbonal-ventral sulcus well im-
pressed. Excurrent siphon truncated, being about 4 the length of the incurrent siphon.

Description. Shell globose, small, reaching 8.5 mm. (about 2 inch) in length and 8
mm. (about inch) in height, light brown to greenish-brown in color, gaping anteriorly
throughout life and closed posteriorly. Anterior slope beaked, the beak extending at least
half the distance to the ventral margin and truncated at nearly a right angle giving the
shell a teredo-like appearance. Beak sculptured with numerous rows of finely denticulated
ridges which parallel the ventral margin. Anterior portion of the disc also sculptured with
close-set rows of denticulated ridges which parallel the anterior margin. Umbonal-ventral
sulcus broad, moderately impressed and sculptured with concentric ridges and growth
lines. Posterior slope tapering to a rounded posterior margin and sculptured with growth
lines. Umbos prominent and located near the anterior third of the shell. Umbonal refiec-
tions narrow, only partially covering the umbos, appressed near the umbos and free an-
teriorly. Dorsal plate consisting of a large, divided mesoplax, each half being ear-shaped
and sculptured with concentric growth lines. Interior of the shell white and glazed. Muscle
scars well marked. Posterior adductor muscle scar large, smooth, elongate-oval in outline,
and bounded on its anterior margin by a shelf. Anterior adductor muscle scar covering
the umbonal reflection. A ventral adductor muscle scar is evident in some specimens. Pal-
lial sinus usually not apparent. Umbonal-ventral sulcus expressed internally as a pro-
nounced ridge with a condyle at its ventral margin. A small chondrophore is present in
the left valve. Internal ligament present but small. Apophyses lacking.

Siphons extending about twice the length of the shell. Aperture of the incurrent siphon
with numerous small cirri. Excurrent siphon truncated, being about one third the length
of the incurrent siphon. A portion of the side walls of the excurrent siphon remains and
this is fringed with a series of lappets which extend from the opening of the excurrent

Plate 89. Xylophaga globosa Sowerby. Fig. 1. Side view of a complete specimen showing the position of the
mesoplax, the pronounced umbonal-ventral sulcus and the truncate excurrent siphon, with only a row of
lappets extending posteriorly on either side of the dorsal surface of the incurrent siphon. This condition is
like that found in X. dorsalis Turton. Fig. 2. Internal view of valve showing the chondrophore, the posi-
tion of the posterior adductor muscle scar, the shelf and the pronounced ventral condyle. Fig. 3. Dorsal
view of shell showing the mesoplax in place. Fig. 4. Dorsal view of mesoplax. Fig. 5. Ventral view of
mesoplax. Fig. 6. Side view of mesoplax. All from Piedra Azul, 7 miles east of Puerto Montt, Chile.

150 JOHNSONIA, No. 34 Xylophaga

siphon along the dorsal surface of the incurrent siphon to its posterior extremity. Man-
tle and siphons light ivory in color and smooth. Foot large, diamond shaped and truncate.
The above description of the soft parts is based on preserved material.

length height ratio h:]
8.5 mm. 8 mm. 1.06 About 7 miles east of Puerto Montt, Chile
8. 0 7 ] : ] ee ee oe se ee

Types. The type of Yylophaga globosa Sowerby is in the British Museum; the type
locality is Valparaiso, Chile in waterlogged wood dredged in 60 fathoms, Hugh Cuming
collector.

Remarks. Xylophaga globosa is closely related to_X. dorsalis Turton. It differs in the
elongate-oval shape of the posterior adductor muscle scar, the less inflated posterior slope,
the larger beak which often extends more than half the distance to the ventral margin,
and the more pronounced umbonal-ventral sulcus. In addition, the shape of the meso-
plax is quite different as shown in the figures for the two species. The excurrent siphon
of XY. dorsalis is truncated at the posterior margin of the valve while that of elobosa ex-
tends to about one third the length of the incurrent siphon.

Xylophaga globosa Sowerby, while not common, is probably not an excessively rare
species even though there are very few specimens in museum collections. The University
of Lund Expedition to Chile collected this species at five stations near Puerto Montt,
Chile and at two of these it was fairly abundant. The specimens were usually taken from
submerged waterlogged wood dredged in 6 to 185 fathoms. One lot was obtained from
a test board submerged in Puerto Montt Harbor in 8 meters. Apparently the species is
not present in sufficient numbers to be of any great economic importance at this locality.
In the samples examined the specimens had bored into the wood to a depth only slightly
greater than the length of the shell, and usually across the grain.

Range. From Panama (Dall 1909) south to Chiloe Island, Chile.

Specimens examined. CHILE: Puerto Montt Harbor (S. Lat. 41°29/10’; W. Long.
72°57'47') in 12 fathoms; Tenglo Canal, Seno Reloncavi, near Puerto Montt (S. Lat.
41°29/33/’; W. Long. 72°58’88”) in 6 fathoms; Bay of Puerto Montt between Isla Ten-
glo and Punta Pilluco (S. Lat. 41°80/05’""; W. Long. 72°56/22”) in 135 fathoms; Piedra
Azul, Seno Reloneavi, about 7 miles east of Puerto Montt (S. Lat. 41°31/30”; W. Long.
72°48/15'’) in about 30 fathoms; Canal San Antonio, Golfo de Ancud (S. Lat. 41°44/10”;
W. Long. 73°15/15”’) in 8 fathoms. All Chiloe Province (all University of Lund Expe-
dition to Chile).

Xylophaga mexicana Dail
Plate 90

Xylophaga mexicana Dall 1908, Bulletin Museum of Comparative Zodlogy 43, no. 6, p. 425 (Albatross, sta-
tion $422, off Acapulco, Mexico in 141 fathoms).

Distinctive characters. Beak extending two thirds the distance to the ventral margin.
Umbonal reflection narrow, slightly recurved and partially covering the umbo. Umbonal-
ventral sulcus narrow, deep and with a small ridge at the posterior margin. Posterior
adductor muscle sear large, oval, smooth and only lightly impressed.

Eastern Pacific JOHNSONIA, No. 34 151

Description. Shell globose, small, reaching 5 mm. (about 4 inch) in length and 4.2 mm.
(about 4 inch) in height. Beaked and widely gaping anteriorly throughout life, rounded
and closed posteriorly. Beak large, extending about two thirds the distance to the ven-
tral margin. Beak sculptured with numerous rows of finely denticulated ridges which are
parallel to the ventral margin. Anterior portion of the dise sculptured with crowded rows
of finely denticulated ridges which are somewhat coarser than those of the beak. Umbonal-
ventral sulcus rather narrow and deep, with a definite ridge at the posterior margin. Pos-
terior slope inflated and broadly rounded, sculptured only by growth lines. Umbos
prominent and located near the anterior third of the shell. Umbonal reflection narrow,
free and only slightly recurved anteriorly. Posteriorly it is appressed but covers only the
anterior half of the umbos. Dorsal plates unknown. Interior of the shell white and glazed.
Muscle scars barely visible on the limited material available for study. Posterior adduc-
tor muscle scar large, oval in outline, smooth and only lightly impressed. It is located
high on the posterior slope. Anterior adductor muscle scar covering most of the umbonal
reflection. Umbonal-ventral sulcus expressed internally as a narrow, high, transversely
grooved ridge, the grooves giving the ridge a segmented appearance. Chondrophore
small. Apophyses lacking.

length height ratio h:]
5 mm. 4.2 mm. i Holotype

Types. The holotype of Xylophaga mexicana Dall is in the United States National
Museum, no. 122947, from the Albatross, station 3422, off Acapulco, Mexico (N. Lat.
16°47’: W. Long. 99°59’) in 141 fathoms.

Remarks. Though this species is based on a single, poor, dead specimen, it appears to
be quite distinctive. Lacking the dorsal plate, it is impossible to say definitely where this
species belongs. However, the size of the beaks, the nearly smooth muscle scars, the
broadly reflected dorsal margin and the deep, narrow umbonal-ventral sulcus with a pro-
nounced ridge at its posterior margin relate it most closely to _Y. globosa Sowerby. In
fact it may prove to be this species.

Range. Known only from the type locality.

Plate 90. Xylophaga mexicana Dall. Fig. 1. External view of the valve of the holotype showing the pro-
nounced ridge on the posterior margin of the umbonal-ventral suleus, and the beak which extends more
than half the distance to the ventral margin. Fig. 2. Internal view of the valve of the holotype showing
the umbonal-ventral ridge and the muscle scar. From off Acapulco, Mexico.

152 JOHNSONIA, No. 34 Xylophaga

Specimens examined. Mexico: Albatross, station 3422, off Acapulco, Mexico (N. Lat.
16°47’; W. Long. 99°59’) in 141 fathoms.

Xylophaga atlantica Richards
Plates 4 and 91

Xylophaga atlantica Richards 1942, Nautilus 56, p. 68, pl. 6, fig. 4 (east coast of the United States).

Distinctive characters. Beak extending less than half the distance to the ventral mar-
gin. Mesoplax triangular, divided and located anterior to the umbos. Umbonal-ventral
suleus shallow with a median thread-like groove. Posterior adductor muscle sear elongate-
oval and irregularly marked. Excurrent siphon nearly as long as the incurrent siphon.

Description. Shell globose, small, reaching 14.5 mm. (about 4 inch) in length and 15
mm. (about $ inch) in height, white to light brown in color and fragile. Gaping anteri-
orly throughout life and closed posteriorly. Anterior slope beaked, the beak extending
less than half the distance to the ventral margin and truncated at nearly a right angle,
giving the shell a teredo-like appearance. Beak sculptured with numerous rows of den-
ticulated ridges which parallel the ventral margin of the beak. Anterior portion of the
dise sculptured with denticulated ridges paralleling the anterior margin. The ridges of the
dise are more closely spaced and coarsely denticulated than those of the beak. Umbonal-
ventral sulcus shallow, with a median thread-like groove. Posterior slope inflated and
having a high, ear-like lobe posteriorly. It is smooth and sculptured only with faint growth
lines. Umbos prominent and located near the anterior third of the shell. Dorsal margin
of the valves reflected anterior to the umbos only. Reflection narrow and free. Mesoplax
minute, divided and located anterior to the umbos. The two parts are triangular in out-
line and have a small cavity below into which the posterior portion of the anterior ad-
ductor muscle extends. Mesoplax of young similar to that of the adult. Interior of shell
white and glazed. Muscle scars well marked. Posterior adductor muscle scar large, oval
in outline, located high on the posterior slope and irregularly marked. Anterior adduc-
tor muscle scar covering most of the reflection of the valves. Pallial sinus not apparent.
Umbonal-ventral sulcus expressed internally as a pronounced ridge with a condyle at its
ventral margin. A rather large chondrophore is present in the left valve. Internal liga-
ment clearly visible. Apophyses lacking. Siphons white to light ivory in color, smooth,
devoid of periostracum and probably not extending more than the length of the shell.
Siphons united, the excurrent siphon being about three fourths the length of the incur-
rent siphon. Opening of the excurrent siphon surrounded by 6 to 8 large papillae. Aper-
ture of incurrent siphon with numerous small papillae. Foot and mantle white. Foot
nearly circular in outline and truncate.

length height ratio h:]

14.5 mm. 15.0 mm. .96 off Cape Breton, Nova Scotia
11.0 10.5 1.04 Holotype
9.5 9.5 1.00 off Block Island, Rhode Island
8.0 eo 1.06 Mount Desert Island, Maine
8.0 8.5 .94 Perry, Maine

Types. The holotype of Xylophaga atlantica Richards is in the Academy of Natural
Sciences Philadelphia, no. 178741. The locality as given by Richards was the east coast

Western Allantic JOHNSONIA, No. 34 153

of the United States. The type locality is here restricted to Mount Desert Island, Maine,
a locality from which specimens are known to have been taken from fixed structures.

Remarks. It is surprising that this Western Atlantic species remained undescribed
until 1942. All early references to Xylophaga dorsalis Turton along our Atlantic coast
are actually to this species, as_X. dorsalis of Europe is not found in our waters. The shells
of the two species are superficially similar but Y. atlantica is a much larger species with
a small mesoplax which is located anterior to, rather than over, the umbos. The posterior
adductor scar of X. at/antica is elongate-oval in outline and is marked with irregular
grooves, while that of XY. dorsalis is more or less circular in outline and nearly smooth.
In addition, the excurrent siphon of _X. at/antica is nearly as long as the incurrent siphon
while in X. dorsalis it is truncated at the posterior margin of the valve.

Xylophaga atlantica Richards is most closely related to_Y. washingtona Bartsch of the
Eastern Pacific but may be separated from that species by its irregularly marked posterior
adductor muscle scar and its broadly triangular mesoplax which has a very small ventral
portion. See also remarks under _X. washingtona Bartsch.

Very little is known concerning the biology of this species. Like others in the genus
Xylophaga it appears to be essentially pelagic. It is seldom taken in fixed structures,
most specimens coming from floating or waterlogged wood. It ranges from off Quebec
south to Virginia and has been taken alive from waterlogged wood brought up from
depths of 50 fathoms. There are numerous records of specimens taken from wood dredged
in 100 to over 1000 fathoms but no mention was made as to whether the specimens were
living or dead. Under Specimens examined the fathoms for each of the U.S. Fisheries
stations is given but this does not necessarily indicate that the specimens were taken alive
at such depths.

The young of Xylophaga atlantica Richards are held within the brood pouch until the

Plate 91. Xylophaga atlantica Richards. Fig. 1. Internal view of left valve showing the irregularly sculptured
posterior adductor muscle scar. From off Cape Breton Island, Nova Scotia. Fig. 2. External view of the
left valve of the holotype showing the shallow umbonal-ventral sulcus with the thread-like median groove.
Fig. 3. Dorsal view of shell showing the mesoplax in place. Fig. 4. Dorsal view of mesoplax. Fig. 5.
Ventral view of mesoplax. Fig. 6. Siphons of a preserved specimen showing that the excurrent siphon is
nearly as long as the incurrent siphon. All from Perry, Maine.

154 JOHNSONIA, No. 34 Xylophaga

late veliger stage. The embryonic shell is about $ mm. in length with pronounced con-
centric sculpture, simple, straight hinge line and prominent umbos. Metamorphosis of
the shell into the adult form begins even before the young are released. The embryonic
shell is figured on plate 4.

Range. From St. Lawrence estuary, Quebec, Canada south to Cape Henry, Virginia.

Specimens examined. CANADA: Trois Pistoles, St. Lawrence Estuary, Quebec (USNM);
St. Pierre Bank, about 180 miles east of Cape Breton Island; about 270 miles southeast
of Cape Breton Island, in atrawl marker buoy (both J. Miller). Marne: Perry; Mount
Desert Island (both MCZ): off North Haven in 10 fathoms (USNM). Massacuuserts:
Rockport (MCZ):; Annisquam River, Gloucester (J. Miller); 10 miles east of Boston
Lightship (MCZ): Brown’s Bank, about 260 miles east of Boston(USNM); off Cohas-
set (J. Miller); Provincetown (MCZ); Speedwell, station 194, 180 miles east of Gloucester
(N. Lat. 42°33’: W. Long. 69°58’) in 110 fathoms; Speedwell, station 198, about 120
miles east of Manchester (N. Lat. 42°30’; W. Long. 70°20’) in 50 fathoms; Speedwell,
station 234, about 60 miles east of Salem (N. Lat. 42°30’; W. Long. 70°38’) (all USNM);
Georges Bank, 155 miles east of Cape Cod (N. Lat. 41°20’; W. Long. 66°50’) (MCZ);
Albatross, station 2076, off Georges Bank, 180 miles east of Cape Cod (N. Lat. 41°13’;
W. Long. 66°00’), in 906 fathoms (USNM); Georges Bank, 65 miles east of Cape Cod
(N. Lat. 14°18’; W. Long. 68°40’) in 50 fathoms (MCZ). Ruove IsLanp: off Block
Island (J. Miller). New Jersey: Mish Hawk, station 1162, about 190 miles east of Sandy
Hook (N. Lat. 40°32’; W. Long. 70°39’) in 45 fathoms; Albatross, station 2550, 160
miles east of Barnegat Bay (N. Lat. 89°44’; W. Long, 70°30’); Fish Hawk, station 998,
120 miles east of Barnegat Bay (N. Lat. 39°43’; W. Long. 71°32’) in 802 fathoms; Fish
Hawk, station 1140, 105 miles east of Atlantic City (N. Lat. 89°34’; W. Long. 71°56’)
in 874 fathoms; Albatross, station 2041, 285 miles due east of Atlantic City (N. Lat.
39°22’; W. Long. 68°25’) in 1608 fathoms; Albatross, station 2036, about 270 miles east
of Cape May (N. Lat. 88°52’; W. Long. 69°24’) in 1735 fathoms (all USNM). Vireinia:
Albatross, station 2736, Hampton Roads in 11 fathoms; A/dbatross, station 2566, 405
miles east of Cape Charles (N. Lat. 87°23’; W. Long. 68°08’) on surface; Albatross, sta-
tion 2731, about 75 miles east southeast of Cape Henry (N. Lat. 36°45’; W. Long. 74°28’)
in 781 fathoms; A/batross, station 2012, 65 miles east southeast of Cape Henry (N. Lat.
36°41’; W. Long. 74°39’) in 66 fathoms; Albatross, station 2729, 75 miles east southest
of Cape Henry (N. Lat. 36°36’; W. Long. 74°32’) in 679 fathoms (all USNM).

Xylophaga washingtona Bartsch
Plate 92
Xylophaga washingtona Bartsch 1921, Proceedings Biological Society Washington 34, p. 32 (San Juan Is-
land, Washington).
Xyloghaga californica Bartsch 1921, Proceedings Biological Society Washington 34, p. 32 (off Pt. Pinos
Light, California).

Distinctive characters. Beak extending less than half the distance to the ventral margin.
Umbonal-ventral sulcus broad and shallow. Mesoplax triangular in outline, divided and
located anterior to the umbos. Posterior adductor muscle scar elongate-oval in outline
and regularly marked with a central and radiating grooves.

> i

Sas” |

Eastern Pacific JOHNSONIA, No. 34 155

Description. Shell globose, small, reaching 6 mm. (about 4 inch) in length and 6.3
mm. (about 4 inch) in height. Widely gaping anteriorly throughout life and closed pos-
teriorly. Anterior slope beaked, the beak extending less than half the distance to the
ventral margin and truncated at nearly a right angle giving the shell a teredo-like appear-
ance. Beak sculptured with numerous rows of finely denticulated ridges, the rows run-
ning parallel to the ventral margin. Anterior portion of the disc sculptured with finely
denticulated ridges which parallel the anterior margin. The ridges of the disc are more
closely spaced and more coarsely denticulate than those of the beak. Umbonal-ventral
sulcus wide and shallow, and sculptured only by faint growth lines. Posterior slope in-
flated, rounded posteriorly, with a high ear-like lobe, and sculptured only by growth lines.
Umbos prominent and located near the anterior third of the shell. Dorsal margin of the
valves reflected anterior to the umbos only. Reflection narrow and free. Dorsal plate
consisting of a minute, divided, triangular mesoplax which is located anterior to the um-
bos and partially encloses the anterior adductor muscle. Interior of the shell white and
glazed. Muscle scars well marked. Posterior adductor muscle scar large, oval in outline
and located high on the posterior slope. The scar is rough, but regularly marked with a
central groove and numerous radiating grooves. Anterior adductor muscle scar covering
the umbonal reflection. Pallial sinus not apparent. Umbonal-ventral sulcus expressed
internally as anarrow and high ridge with a condyle at its ventral margin. A rather large
chondrophore is located on the left valve. Internal ligament present. Apophyses lacking.

length height ratio h:]
6.0mm. 6.3mm. 0.95 Station 2867, off coast of Washington
5.9 6.0 0.98 ee ee eé se ee ee
5.8 5.2 iJ Holotype, Xylophaga washingtona Bartsch
4.8 4.8 1.0 Holotype, Xylophaga californica Bartsch

Types. The holotype of Xylophaga washingtona Bartsch from San Juan Island, Wash-
ington is in the United States National Museum, no. 334479, as is that of XY. californica
Bartsch, no. 209876, from Albatross, station 4523, off Point Pinos Light, California.

Plate 92. Xylophaga washingtona Bartsch. Fig. 1. Dorsal view of paratype showing the mesoplax in place.
Fig. 2. External view of right valve of the holotype showing the shallow, smooth umbonal-ventral sulcus.
Fig. 3. Internal view of left valve of the holotype showing the regularly marked posterior adductor mus-
cle scar. Fig. 4. Ventral view of mesoplax. Fig. 5. Dorsal view of mesoplax. Fig. 6. Side view of meso-
plax. All specimens from San Juan Island, Washington.

156 JOHNSONIA, No. 34 Xylophaga

Remarks. Xylophaga washingtona Bartsch appears to be most closely related to Xylo-
phaga atlantica Richards of the Western Atlantic. It is, however, a smaller species, the
mesoplax is longer in proportion to its width, with a larger ventral portion, and the pos-
terior adductor muscle sear is regularly marked. In addition, the umbonal-ventral sulcus
of XY. atlantica Richards has a distinct, thread-like median groove which corresponds to
the internal ridge. From Xylophaga mexicana Dall this species differs by having the
beak extending less than half the distance to the ventral margin and by having a shallow,
broad umbonal-ventral sulcus, not bounded by a distinct posterior ridge. In addition,
the posterior adductor muscle scar is large and distinctly marked.

The type specimens of Yylophaga californica Bartsch are small, poor specimens of _X.
washingtona Bartsch. Both species were described in the same paper and though XY. ca/l-
ifornica was the first described, the specimens were so poor and fragmentary that it
seemed best to place it in the synonymy of Y. washingtona of which Bartsch had good
and ample material.

Specimens examined. WASHINGTON: Friday Harbor, San Juan Islands (MCZ); Alba-
tross, station 3456, Straits of Juan de Fuca (N. Lat. 48°31’; W. Long. 124°43’) in 136
fathoms (USNM); Ballard Bay in 10 fathoms(W. J. Eyerdam):; Departure Bay ; A/ba-
tross, Station 2867, off Flattery Rocks (N. Lat. 48°07’: W. Long. 124°55’) in 37 fathoms
(both USNM). Orecon: Albatross, station 5432, about 20 miles off Cape Meares Light-
ship in 97 fathoms (USNM). Catirornia: Albatross, station 4523, off Point Pinos Light
in 75-108 fathoms (USNM).

Xylophaga abyssorum Dail
Plate 98

Xylophaga abyssorum Dall 1886, Bulletin Museum Comparative Zodlogy 12, p. 317, pl. 9, figs. 7, 7a (Blake,
station 215, N. Lat. 13°51’; W. Long. 61°03’, off St. Lucia in 226 fathoms).

Distinctive characters. Beak extending less than half the distance to the ventral mar-
gin. Umbonal-ventral sulcus shallow and bounded on its posterior margin by a pro-
nounced keel. Posterior adductor muscle scar broadly oval and irregularly roughened.

Description. Shell minute, globose, reaching 4 mm. (about + inch) in length and 3.5
mm. (about 4 inch) in height, white in color and fragile. Beaked and widely gaping an-
teriorly throughout life, rounded and closed posteriorly. Beak extending less than half
the distance to the ventral margin and truncated nearly at a right angle to the anterior
margin of the dise, giving the shell a teredo-like appearance. Beak sculptured with numer-
ous rows of finely denticulated ridges, the rows running parallel to the ventral margin of
the beaks. Anterior portion of the disc sculptured with crowded rows of denticulated
ridges which run parallel to the anterior margin of the disc. The ridges of the disc are
more closely crowded and more coarsely denticulated than those of the beaks. Remainder
of the disc composed of a broad, rather shallow umbonal-ventral sulcus which is sculp-
tured only by growth lines. Ridge bounding the anterior edge of the sulcus, low and
rounded, the one on the posterior edge high and sharp. Posterior slope inflated, rounded
and sculptured with concentric growth lines and low ridges. Umbo prominent, located
near the anterior third of the shell. Dorsal margin of the valves reflected anterior to the
umbos only. Accessory plates unknown. Interior of shell white and glazed. Muscle scars

Western Atlantic JOHNSONIA, No. 34 157

well marked. Posterior adductor muscle scar large, broadly oval in outline and roughened.
Anterior adductor muscle scar covering a small portion of the dorsal reflection. Pallial
sinus not apparent. Umbonal-ventral sulcus expressed internally as a low, beaded ridge
with a large condyle at the ventral margin. A small chondrophore is present in the left
valve. Apophyses lacking.

length height ratio h:]
4.8mm. 4.5 mm. 1.0 off Atlantic City, New Jersey
4.0 3.5 fel Holotype
2.0 ONO, 1.0 off Cape Hatteras, North Carolina

Types. The holotype of Xylophaga abyssorum Dall is in the Museum of Comparative
Zoblogy, no. 8135. The type locality is Blake, station 215 (N. Lat. 18°51'; W. Long.
61°03’), off St. Lucia, Lesser Antilles in 226 fathoms.

Remarks. This rare species is easily distinguished from others in the genus by its mi-
nute size and by the strong keel which separates the disc from the posterior slope. Dall
in his original description stated that he had not seen any accessory plates nor an attach-
ment area for them. However, asingle specimen from off Cape Hatteras, North Carolina
had a fragment of a plate remaining but this was too incomplete to attempt a description.

It would appear from the record that this species may bore into mud, sand and soft
stone. This habit is quite contrary to that of all other Yylophaga. However, all specimens
studied were dredged from considerable depths and all appeared to have been dead at the
time they were taken. Consequently, little can be said as to their true habitat.

Range. From off Atlantic City, New Jersey south through the Lesser Antilles to
St. Lucia.

Specimens examined. NEw JERsEy: Albatross, station 2095 (N. Lat. 39°29’; W. Long.
70°58’) about 200 miles off Atlantic City in 1342 fathoms in globigerine ooze (USNM).
Nort Carouina: Albatross, station 2115 (N. Lat. 35°49’; W. Long. 74°34’) about 40
miles off Cape Hatteras in 843 fathoms in mud and fine sand (USNM). Fiorina: Al
batross, station 2668 (N. Lat. 30°58’; W. Long. 79°38’) about 68 miles off Fernandina
in 294 fathoms (USNM). Lesser AnrILLEs: Blake, station 192 (N. Lat. 15°17’; W.
Long. 61°24’) off Dominica in 138 fathoms (USNM); Blake, station 215 (N. Lat. 18°51’;
W. Long. 61°03’) off St. Lucia in 226 fathoms in a coral nodule (MCZ).

Plate 93. Xylophaga abyssorum Dall. Fig. 1. External view of the left valve of the holotype. Fig. 2. Dorsal
view of the holotype showing the pronounced ridge on the posterior margin of the umbonal-ventral sulcus
(Fig. 1 and 2 after Dall 1886). Fig. 3. Internal view of the right valve of the holotype to show the large
umbonal-ventral ridge and the muscles scars.

158 JOHNSONIA, No. 34 Xylophaga

Key for the identification of the genera of the Pholadidae

(based upon complete adult specimens)

1. Adult shell with a callum . sagt j
Adult shell without a callum Se hit es 5 ie ee ww
9... ‘Shell. with apophyses. <A: 2.4. 4" -) aetna oe ee
Shell without apophyses ... . Ree a re eg er
3. Shell with more than one dorsal See ener E st pee ee a
Shell with-only one-dorsal plate: . 7.0 24. 2 Sen ee ee ee
A Shell with protoplaxvomllys.. 28 a) ren eee eens ee S IC
Shell with mesoplaxonly=. . < «=. 5 \ KARL: & Pea amb

Shell beaked anteriorly, mesoplax in one piece, aceeee ena sulcus
present... 8.0: ee TM ee ee a een A
Shell rounded ean: mesoplax in two pieces, umbonal-ventral sulcus

Or

not present"... .'. Be aca pe ee a ee . «a 2 Falona
6. Shell with a largely chatewees protoplax, a fieioaix but no metaplax,
umbonal reflection not septate. . ....... .. . .. . Cyrtopleura
Shell with a calcareous protoplax, mesoplax and metaplax, umbonal re-
flection: septate. +a eo ae ee ee, de Se ee
7. Shell with.apoplyseso {c: 4p" et eee Bye ee. eee cee ee ee
Shell without apophyses ... . : 3 met eeeaee. eS
8. Animal capable of complete retraction eithin ne shell ohne cage ree
Animal not capable of complete retraction within the shell. . Chaceia
§,- Dorsal’ plate ‘one, a mesoplax“; 04 4. < . a 2 SS ee
Dorsal plates two, a mesoplax and a metaplax .. . ~~.) 2-: os 42
1O.° Mesoplax divided longitudinally =, ......... 272". fed).e. sees
Mesoplax not divided longitudinally . . . . a, es «= Pentiella

11. Siphonoplax present, callum extended dorsally nears the beaks. Rock
borersm-. sae wo dw. 3 Slee tale. oe ee ee ere ers
Siphonoplax nas callum not extending dorsally beyond the beaks.
Wood borers . . . : alee Pad & eee eeu cen) er lapis

12. Valves divided into ieee aisaaet areas, posterior area with overlapping
chitinous plates. Shell usually one or more inches in length Parapholas
Valves divided into two areas, posterior area without chitinous plates
Shell usually under one inch in length . .. . a eee. ses

13. Shell with funnel-shaped pit below the umbonal Sarena: callum not
extending between the beaks. Wood borers . . . . . . . . Martesia
Shell with the umbonal reflection closely appressed. Callum extending
between the beaks and on either side of the mesoplax. Shell and rock
borers. 4._ 3 cis Seas ok dhe a ; vow 2 & . Diplathyra

14. Shell with soriplces nal overlapping eee fevoneria developed on
right valve only... . A, : : oi» Sa Jomannetia
Shell with incomplete sated Senon age dec on both valves
equally 9 2 (i g.hd, ee et a ae ee eee ae Vereen a

‘An African genus, not included in this report.

Notes
Genus Penitella Valenciennes

Unfortunately the genus Navea Gray was omitted from the synonymy of Penitella
which was considered on page 70. The name was based upon the young stage of a Penitella.
Navea Gray 1851, Annals and Magazine of Natural History (2) 8, p. 335 (genotype, N. subglobosa Gray,

subsequent designation Stoliczka 1870).

Western Atlantic JOHNSONIA, No. 34 159

Barnea maritima Da//

Barnea maritima ‘d’Orbigny’ Dall 1889, Bulletin United States National Museum, no. 37, p. 72 (Texas)
[nomen nudum].

It has been impossible to find any record of this name in d’Orbigny’s writings. It may
possibly have been a manuscript name on a specimen in the collection of the United
States National Museum at the time Dall was compiling his list of the marine mollusks
of the Southeastern Coast of the United States. Following Dall, C. W. Johnson included
the name in his ‘‘Marine Mollusca of the Atlantic Coast from Labrador to Texas”’ and
since then it has been mentioned in several other lists dealing with the Texas area though
no one has ever described or figured it.

Pholadidea tridens Gray

Talona tridens Gray 1843 in Dieffenbach’s Travels in New Zealand 2, p. 254 (New Zealand).
Pholas tridens Gray, Sowerby 1849, Thesaurus Conchyliorum 2, p. 10, p. 398, pl. 106, figs. 60-61.
Pholadidea tridens Gray, Dall 1909, Proceedings United States National Museum 37, no. 1704, p. 277.

This species has been recorded by Sowerby, as coming from Montecristi, Ecuador,
Hugh Cuming collector, and Dall (1909) lists it in his catalogue of Peruvian mollusks.
However, we have not seen a specimen from the Eastern Pacific. Dall states (op. cit.,

p. 188) that he has included species ‘‘. . . belonging to a widely distributed group, such as
the Pholadidae, though not actually reported from a Peruvian locality . . . knowing that in

all probability it will be found on more thorough search in Peruvian territory.’* Dall’s
inclusion of P. tridens Gray was undoubtedly based on Sowerby’s record which was based
on Cuming material, a collection known to contain many errors of locality.’ Consequently
it seems best to omit this species from the Eastern Pacific fauna until material with
known locality is obtained.

Scyphomya semicostata Lea

Pholas semicostata H. C. Lea 1844, Proc. Boston Society Natural History 1, p. 204 (South Carolina); Lea
1845, Boston Journal Natural History 5, p. 285, pl. 24, fig. 1.

Zirfaea semicostata Lea, Dall 1889, Bulletin United States National Museum, no. 37, p. 72.

Scyphomya semicostata Lea, Dall 1898, Transactions Wagner Free Institute of Science, Philadelphia 3, pt.
4, p. 822.

This species was described from a single, minute specimen in poor condition and with-
out definite locality. Even the locality, South Carolina, is open to question as Lea (1845)
states ‘‘. . . this curious little species I found among some shells sent to my father many
years since, from South Carolina.’” The species has been shifted from one genus to
another and in 1898 Dall created the genus Scyphomya for its reception. At the time
Dall created the genus he had not seen Lea’s specimen, in fact, the location of the type
is unknown. Specimens in the United States National Museum labeled by Dall as sem-
costata Lea proved to be Jouannetia quillingi Turner. However, it is not certain that
these were the specimens on which Dall based his generic name. Species in the genus
Jouannetia lack apophyses but semicostata Lea, the type of Scyphomya, has them. Dall
mentions the apophyses in the original description of his genus but states that they
were missing in his specimens. Until more data are at hand it is impossible to place
properly this genus and species.

'Clench 1945, Occasional Papers On Mollusks 1, p. 26.

160 JOHNSONIA, No. 34 References Cited

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Harvey, E. N. 1952, Bioluminescence. Academic Press Inc., New York, pp. 1-649, 187 text figures.

Jeffreys, J. G. 1865, British Conchology, London 3, pp. 93-122.

Johnson, C. W. 1904, Nautilus 18, pp. 100-103, 1 text figure.

Johnston, G. 1850, An Introduction to Conchology, London, pp. 149-162.

Kellogg, J. L. 1915, Journal of Morphology 26, pp. 625-701.

Kuhnelt, W. 1942, Palaeobiologia 7, pt. 5-6, pp. 428-447.

Lamy, E. 1921, Bulletin Muséum National d’Histoire Naturelle 27, pp. 178-183.

Lamy, E. 1922, Bulletin Muséum National d’ Histoire Naturelle 28, pp. 243-246.

Lamy, E. 1925-1926, Journal de Conchyliologie 69, pp. 19-51, 79-103, 136-168, 193-222.

Lebour, M. V. 1938, Journal Marine Biological Association United Kingdom 23, pp. 119-144.

Lloyd, F. E. 1897, Transactions New York Academy Sciences 16, pp. 307-316, pls. 25-26.

MacGinitie, G. E. 1935, American Midland Naturalist 16, pp. 629-765.

Moore, D. D. 1947, Port of Sydney Journal 1, no. 3, pp. 74-75.

Morch, O. A. L. 1876, Journal de Conchyliologie 24, p. 367.

Okada, Y. K. 1927, Bulletin Société Zoologique de France 52, pp. 95-98.

Poli, J. X. 1791, Testacea Utriusque Siciliae Eorumque Historia et Anatome 1, pp. 39-50, pls. 7-8.

Purchon, R. D. 1941, Journal Marine Biological Association United Kingdom 25, pp. 1-39, 16 text figures.

Ridewood, W. G. 1903, Philosophical Transactions Royal Society London (B) 195, pp. 147-284.

Sieverts, H. 1933, Neues Jahrbuch fur Mineralogie, Geologie und Palaontologie, Beilage-Band 71, pp. 267-
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Sigerfoos, C. P. 1895, Annales Magazine Natural History (6) 16, pp. 233-238.

Smith, J. 1894, Proceedings and Transactions Natural History Society, Glasgow (N.S.) 4, pp. 37-39.

Sowerby, G. B. 1849, Monograph of the Genus Pholas. Thesaurus Conchyliorum 2, pt. 10, pp. 485-505, pls.
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JOHNSONIA

Published by
THe DerparTMENT oF Mo.Luusks
Museum of Comparative Zodlogy, Harvard University

Cambridge, Massachusetts

BAdYAND

URVESSITY

VOL. 3. NO. 35

JANUARY 30, 1956 MELONGENIDAE

THE FAMILY MELONGENIDAE IN THE WESTERN ATLANTIC
BY
WILLIAM J. CLENCH AND Ruri ID. TURNER

The family Melongenidae possesses only a few species, but these occur in nearly all
tropical and temperate seas. All are intertidal or live a little below low water and, so far
as is known, are generally predatory in habit though occasionally feeding on dead animal
matter. In certain areas, such as western Florida, they may be sufficiently abundant on
oyster bars to be a rather serious menace.

Melongena has presented many challenging problems. This is particularly true of the
polytypic species complex of Melongena corona Gmelin. Populations of this species only
short distances apart may be strikingly different. Certain populations are very uniform
while others show an amazing amount of variation (Plate 100). This type of variation
suggests that some factor in their life cycle prevents wide dispersal of the young, and that
the progeny of a colony remains within a limited area. Marine species having young
which are readily dispersed are usually either quite uniform throughout their range or

Plate 94. Veliger larvae of Melongena corona Gmelin, from Punta Rassa, Florida. Fig. 1. Late veliger stage
(27 days old) just before emerging from the capsule, showing the four velar lobes, the tentacles, the eyes
and the shell. Fig. 2. A specimen at the same stage with the animal partially retracted within the shell.
The well-developed foot can be seen in the center with the velar lobes behind. The shell at this stage has
one and one half whorls. (about 75x)

161

162 JOHNSONIA, No. 35 Melongena

else show a gradient of change or a cline from one end of the range to the other. In order
to determine the controlling factor, we kept specimens in an aquarium through the
breeding season and found that all of the larval stages are passed within the egg capsule,
the emerging young crawling snails being capable of only limited dispersal. Such young
snails certainly could reach other colonies by mechanical means and this no doubt occa-
sionally happens. However, mechanical transport on floating mangrove leaves or other
debris appears to be rather uncommon and as a consequence gene flow between colonies
is probably limited.

Early in March 1952, Dr. and Mrs. David Schmidt sent us by air mail six live speci-
mens of Melongena corona corona Gmelin from Punta Rassa, Florida. The following
notes on the egg laying and development are based on these specimens. ‘The eggs were
deposited in light, straw-yellow capsules about 12 mm. in diameter. The capsules were
composed of layers of tough, striated, chitinous material and were deposited in strings.
Under natural conditions they are attached to rocks, shells or other hard surfaces.

On July 11 the first egg capsules appeared. The temperature of the water at that time
was 78° EF. ‘'wo females observed during oviposition deposited the strings of capsules on
the side of the aquarium in the same area. The capsules were attached to a base mem-
brane and the first string was composed of eighteen capsules, the second of fifteen, the
third of nine, and the fourth string of only seven capsules. ‘The number of eggs per cap-

— ii(2

eS

Plate 95. Egg capsules of Melongena corona Gmelin, from Punta Rassa, Florida. Fig. 1. A single egg capsule
showing the larvae within. Fig. 2. Four capsules in normal arrangement viewed from above and somewhat
diagonally. The small ellipse at the top of each capsule is the point at which the capsule eventually rup-
tures to liberate the young crawling snails (about 6x).

Western Atlantic JOHNSONIA, No. 35 163

sule varied from 50 to 120. The eggs were contained in a jelly-like mass and were a light
yellow in color and opaque. The first cleavage occurred two and one half hours after the
capsule had been produced; it was equal and complete. It was impossible to follow the
cleavages beyond the sixteen-cell stage on living material due to the large amount of yolk
and the consequent opacity of the egg. The early trochophore stage had been reached
by the third day and the embryo rotated slowly by means of short, minute cilia. By the
seventh day the proboscis and foot began to show and the external kidneys were well
developed. The next day a well-developed proboscis, foot and velum were evident as
well as the beginning of the tentacles with the eyes at their base. The embryonic heart
was first seen pulsating at this time. The shell at this stage is small and cap-like. By the
sixteenth day the veliger larvae were well developed. They had four large velar lobes
with a double row of strong cilia on their outer edges. ‘The foot and operculum were well
developed and the shell had one and one half whorls. The animals were actually showing
signs of crawling by the seventeenth day. By the twenty-fifth day the young were actively
crawling within the capsule. The velar lobes were still very active, however, probably
functioning as ‘‘respiratory organs’’ to aid in the circulation of the fluid within the cap-
sule and the consequent diffusion of gas through the capsule wall. The gills and siphon
were well developed at this time. Specimens twenty-five days old or older, when removed
from the capsule,no longer protruded the velar lobes but crawled actively and could rapidly
turn over the shell with the aid of the foot. By this time the escape pore in the egg cap-
sule was very thin. The first capsule ruptured on the twenty-seventh day (August 6).
Most of the young snails, now incapable of swimming, immediately crawled to the water’s
edge and appeared to be feeding on the minute organisms along the sides of the tank.
There were no nurse eggs and approximately ninety-five percent of all the eggs developed.

The radulae of Melongena and Pugilina are surprisingly small, but the individual teeth
are strong, with large hooked denticles, making a very effective rasping tool. The radula
is rachiglossate, consisting of three longitudinal rows of teeth—a central and two lateral

QAI
QRA 1 (Wing
eA | ORY

Plate 96. Radulae of Melongena and Pugilina. Fig. 1. Melongena melongena Linné, Cienfuegos, Cuba. Fig. 2.
Melongena melongena Linné, from a young specimen, Puerto Barrios, Guatemala. Fig. 3. Melongena bicolor
Say, Grassy Key, Florida. Fig. 4. Melongena corona Johnstonei Clench and Turner, Little Lagoon, Gulf
Shores, Alabama. Fig. 5. Melongena corona nta Rassa, Florida. Fig. 6. Pugilina morio

Linné, Ministere Bay, Tobago Island, West MUS eSO%!?. 7001

LIBRARY
JAN 30 195

1 mm

HARVARD

164 JOHNSONIA, No. 35 Melongena

rows. The differences between the several species and genera are slight but appear to be
constant. The various types are figured on Plate 96. The formula is $: 4: 4.

The proboscis is long and can be greatly extended when the need occurs, such as when
feeding on oysters or other bivalves. In the laboratory, a three-inch specimen was able
to reach a small piece of fish placed in the bottom of a six-inch test tube. The proboscis
was so greatly extended that it became nearly transparent and it was possible to see the
minute bits of fish carried along toward the esophagus. Wilcox (1897, p. 27) describes
the feeding of Me/ongena corona under natural conditions.

According to Dall (1890, p. 118) Melongena is first recorded from the Eocene in the
Vicksburg Limestone. Dall’s Melongena sculpturata, which he described from the Mio-
cene of Ballast Point, Florida, appears to us to belong to the genus Pugilina, as does M.
antillarum Gabb from Santo Domingo. Melongena corona first appeared in the Caloosa-
hatchee marl of the Lower Pliocene. There appears to be little or no difference between
these Pliocene specimens and those taken in certain areas of southwest Florida at present.

Plate 97. Melongena corona Gmelin feeding on a piece of fish placed at the bottom of a six inch test tube. In
Fig. 1, the head may be seen inside the test tube with the proboscis fully extended to reach the fish. In
Fig. 2, the Melongena having grasped the food has pulled a piece part way up the tube for easier feeding.
This demonstrates graphically how these animals can so effectively feed on clams (reduced nearly one half).

ACKNOWLEDGEMENTS

As with all previously issued numbers in this series, we are deeply indebted to several
persons who have aided in many ways to make this study.possible. To Dr. and Mrs.
David Schmidt of Sarona, Wisconsin, we are grateful for live material from several lo-
calities in Florida. Their repeated sendings made possible the study of the gross embry-
ology of Melongena corona. To Mr. and Mrs. Harry I. Johnstone of Mobile, Alabama,
we are indebted for a long series of Melongena corona johnstonei from Little Lagoon, Gulf
Shores, Alabama, which has established the westernmost record for this genus in the
northern Gulf area. To Mr. John H. Butler of Marathon, Florida, we are grateful for
specimens of Melongena bicolor Say and the notes on their feeding habits. We are also

Western Atlantic JOHNSONIA, No. 35 165

indebted to C. G. Aguayo, A. C. Bippus, John Finlay, Ruth Merrill, H. A. Pilsbry,
J. J. Parodiz, H. A. Rehder and Jeanne S. Schwengel for the loan of material which has
aided greatly in obtaining an understanding of the distribution of the various species.

Genus Melongena Schumacher

Galeodes Riding 1798, Museum Boltenianum, p. 53; non Galeodes Olivier 1791 { Arachnida].
Melongena Schumacher 1817, Essai d’un Nouveau Systéme des Habitations des Vers Testacés, Copenhagen,
p. 64, 212.

Type species, Melongena fasciata Schumacher (= Murex melongena Linné),monotypic.
Z Cc q . }

Shell imperforate, subglobose to elongate and generally solid to massive in structure.
Sculpture variable, ranging from smooth specimens to those having as many as four rows
of shoulder spines and one row of basal spines. Periostracum usually persistent, yellowish-
brown in color and usually sufficiently transparent to show the coloration beneath. Oper-
culum unguiculate with a basal nucleus.

Iredale’ pointed out that Volema Réding had priority over Melongena, but at the same
time, he designated Volema paradisiaca Réding as the type species. This was a fortunate
choice as Volema paradisiaca (= Murex ficus Gmelin) is certainly not congeneric with
Melongena of the Western Atlantic and Eastern Pacific.

Subgenus Melongena Schumacher

Shell capacious, generally massive in structure and with the later whorls gradually en-
veloping the earlier whorls, leaving an irregular and somewhat deeply channeled suture
and an apparently greatly reduced spire. Spines generally short, broad and spike-like,
and usually developed at right angles to the whorl surface with no tendency to recurve
toward the spire.

This subgenus is limited in its distribution to portions of the West Indies, both coasts
of Central America and the northern coasts of South America.

Melongena melongena Linné
Plates 96 and 98

Murex melongena Linné 1758, Systema Naturae, ed. 10, p. 751 (America).

Galeodes melongena Linné, Réding 1798, Museum Boltenianum, p. 53.

Pyrula melongena Lamarck 1816, Encyclopédie Méthodique (table of plates, p. 8), pl. 435, figs. 8a-e; La-
marck 1822, Animaux Sans Vertébres 7, p. 140 (1’Océan des Antilles).

Melongena fasciata Schumacher 1817, Essai d’un Nouveau Systéme des Habitations des Vers Testacés,
Copenhagen, p. 212 (no locality given).

Melongena melongena Linné, Tryon 1881, Manual of Conchology (1) 3, p. 107, pl. 41, figs. 197-198; Bayer
1952, Zoologische Mededelingen 31, p. 268.

Melongena melongena denudata Dollfus 1887, Bull. Société d* Etudes Scientifiques d’Angers (n.s.) 17, p. 56,
pl. 2, fig. 3 (Antilles).

Melongena melongena multispinosa Dollfus 1887, Bull. Société d’Etudes Scientifiques d’Angers (n.s.) 17,
p. 56, pl. 2, fig. 4 (Antilles).

Melongena melongena semispinosa Dollfus 1887, Bull. Société d’Etudes Scientifiques d’ Angers (n.s.) 17, p.
56, pl. 2, fig. 5 (Antilles).

Melongena margaritana Richards 1943, Journal of Paleontology 17, pp. 120-123, figs. 1-2 (Pleistocene fos-
sil, Juan Griego, Margarita Island, Venezuela).

‘Iredale, T. 1917, Proc. Malacological Society London 12, p. 322.

166 JOHNSONIA, No. 35 Melongena

Description. Shell reaching 182 mm. (about 74 inches) in length, solid and usually
spinose. Color white with dark brown to brownish-purple spiral bands of varying widths.
Generally there is a broad band from the whorl shoulder to the periphery which is sepa-
rated by a white area from a much narrower subperipheral band. Below this subperiphe-
ral band there are usually numerous thread-like bands, rarely a broad solid band. Whorls
74, strongly convex, particularly above the periphery. Spire somewhat elevated in young
specimens, but in the adult stage the spire is depressed, occasionally hardly extending
above the top of the body whorl. Aperture subelliptical with a well-developed anal notch
and a broad, flattened and short siphonal canal. Parietal lip usually heavily glazed.
Palatal or outer lip moderately thin and usually strongly crenulate, particularly below
the peripheral area. Columella nearly perpendicular and twisted. Umbilicus nearly al-
ways closed by the extension of the parietal callus. Suture canaliculate, the channel be-
coming fairly broad in large specimens. Occasionally the later whorls are folded over the
earlier whorls. ‘The broadening of the channel is sometimes due to the presence of spines
on the previous whorl that were developed ahead of the area of the anal canal. Sculpture
consisting of spiral rows of strong spines. Usually there are two or three rows above and
one row below the periphery near the base of the shell. Occasional specimens are com-
pletely devoid of spines and others may have but few. Young specimens very frequently
show a rather well-developed sculpture of small spiral threads. The young specimens and
the early whorls of the adult are usually strongly nodulose. As the shell advances in age
the nodulose costae disappear or are resolved into spines. Operculum horny, unguiculate,
rather solid, with the nucleus at the lower end, and sculptured with rather strong con-
centric lines of growth. Lower surface of the operculum with a large muscle scar which
has a smooth shiny margin and a dull rugose central area. Periostracum quite smooth,
dull and having exceedingly fine axial growth lines.

length width *
182 mm. 144mm. _ Bay of Cardenas, Cuba
179 131 Jamaica
97 71 Monte Cristi, Santo Domingo

* not including the spines

Types. It would certainly appear that Linné did not have a specimen of Melongena
melongena in his collection at the time the original description was written. There is,
however, no question as to what species he was dealing with, as his references are mainly
to recognizable figures of this common West Indian species. We here select Bonanni
1684, Recreatio, fig. 186, to be the type figure, for this was Linné’s first reference. As
Linné gave only America in his original reference we here limit the type locality to King-
ston, Jamaica. Hanley (1855, p. 298, [psa Linnaei Conchylia) states that Linné’s reference
to Lister ‘‘was correctly referred to in the review copy” (i.e., the interleaved copy of the
12th edition that Linné used for his corrections and additions) and that Jamaica had been
added to the label of the specimen then in the Linnean Cabinet, a specimen probably
received after the tenth edition had been published. The holotype of MW. margaritana
Richards is in the Academy of Natural Sciences, Philadelphia, no. 14931, from Juan
Griego, Margarita Island, Venezuela, a Pleistocene fossil.

Remarks. Melongena melongena Linné and Melongena patula Broderip and Sowerby
are exceedingly close in their relationship. In general, M/. melongena never reaches the

Western Atlantic JOHNSONIA, No. 35 167

size that is exhibited by M. patula. Also, M. patula is much darker in coloration. M.
melongena may lack spines or have as many as four rows of spines, while M. patuda,
which also may be without spines, never has more than one row, and this is composed
of relatively few spines. There is considerable variation in the number of rows of spines
in M. melongena, even among specimens obtained from a single locality. We have a large
series from Monte Cristi, Santo Domingo, containing specimens which are completely
smooth and others which have up to four rows of rather large spines.

The distribution of this species in the West Indies is rather remarkable. It appears to
be limited to Cuba, the Isle of Pines, Jamaica and Hispaniola. It does not occur in the
Bahamas or in Puerto Rico. In the Lesser Antilles it occurs only in Trinidad and Tobago
which are on the continental shelf of South America. This is true in part for the records
we have from the Bay Islands and Curacao. These island groups are beyond the conti-
nental shelf but they do lie close in, being only a few miles off the coast. A few of the
earlier records are certainly open to question, like those given by Krebs (1864, p. 18) for
the Virgin Islands and Lesser Antilles. Records from the Florida Keys and the Texas
coast are certainly in error (Dall 1889, p. 112). It must be understood that many of the
early published records were based upon guesses as to the locality; once in print, they
were continually copied until challenged or completely disproved. This was a fault of the
times rather than an intent to mislead. Locality data at that time were only of casual
interest, perhaps added as an afterthought to the all important name of the specimen.
Today the name of a specimen is certainly the least important of all data associated with
it. It is the only thing that can be changed without destroying the real factual data as-
sociated with a specimen. Accurate locality data are the only basis upon which ranges
can be established and the ranges of many species considered together outline patterns of
distribution allowing us to define faunal regions.

Melongena melongena Linné occurs usually in places where there is a moderate ad-

Plate 98. Melongena melongena Linné. Figs. 1-3. Hicacos Peninsula, Cardenas, Cuba. Fig. | (3/5x). Fig. 2
Young specimen (natural size). Fig. 3 (2/3x).

168 JOHNSONIA, No. 35 Melongena

mixture of fresh and salt water, such as at the heads of bays where enough freshwater
enters to produce a somewhat brackish condition.

This species was probably used for food by the Cuban Indians as specimens now in the
Museo Poey, Habana, were found in a kitchen midden at Vertientes, Camagiiey, Cuba.
The same is true of Melongena patula, as many specimens were found in some recent
Indian excavations near Balboa, Panama.

The three names given by Dollfus, denudata, multispinosa and semispinosa, apply only
to individual forms of Melongena melongena, all of which are occasionally found in a
single colony. They certainly have no taxonomic value.

Melongena margaritana Richards appears to be nothing but a young specimen of MW.
melongena Linné. Richards compares his species with M. corona minor Sowerby [| =bi-
color Say] from which, of course, it differs. He stresses the lack of spines as perhaps the
most outstanding character of his new form, but the young as well as many adults of
M. melongena often lack spines.

Range. From Tampico, Mexico south and east along the coast of Central and north-
ern South America as far as Dutch Guiana. In the West Indies it occurs only in Cuba,
Isle of Pines, Hispaniola, Jamaica and some islands off the coast of South America.

Specimens examined. CuBpa: Puerto Esperanza; Bahia Honda; Mariel (all Museo
Poey): Bahia de Cabanas (MCZ:; Museo Poey): mouth of Almendares River (M.Jaume):
Batabana6 (MCZ: USNM):; Matanzas Bay (Museo Poey); Cardenas (MCZ: Museo
Poey:; John Finlay): La Isabela (M. Jaume); Cayo Cristo, Sagua la Grande; Cayo
Francés, Caibarién; Rio Damuji, Donastia and La Milpa, Cienfuegos (all MCZ); Nuevi-
tas, Camagiiey; Punta Carenero, Puerto Padre (both M. Jaume); Gibara (J. Weber:
Museo Poey; USNM): El Cafion, Banes Bay (MCZ): Macabi, Banes Bay (A. Quifiones,
Museo Poey); Mouth of Yateras River (MCZ): Rente, Santiago de Cuba (M. Jaume);
Guantanamo Bay; Manzanillo (both Museo Poey). IsLE or Pines: (D. and N. E.
Schmidt). Hispanro.a: Port au Prince and Fort Liberté, Haiti (MCZ; USNM); Monte
Cristiand Puerto Plata, Santo Domingo (both MCZ). Jamaica: Kingston; Black River,
St. Elizabeth; Port Royal (all USNM): Port Antonio; Bogue, Montego Bay (both A.
Bippus). Lesser AnrILLEsS: Alley Creek; Ortoire River, Mayaro; Carenage; and Ma-
tura Bay, all Trinidad (all H.G. Kugler); Tobago (MCZ). Caripnean Istanps: Utilla
Island, Bay Islands (MCZ): Vignon, Curacao (USNM). Mexico: Tampico; 15 miles
north of Tecolutla, Vera Cruz (both T. Pulley); Tuxpam, Vera Cruz (M. Bourgeois):
Campoton, Campeche (MCZ). GuaremMa.a: Livingston (MCZ); Puerto Barrios (Univ.
of Michigan; USNM): Punta Manabique (USN M). Honpuras: Trujillo(MCZ); Puerto
Cortez; Tela (both USNM). Panama: Colén(USNM). Cotompia: near Santa Marta;
Sabanilla (both USNM); Cartagena (USNM; MCZ); Puerto Colombia (USNM).
VENEZUELA: Isla de Margarita (USNM). Brirish Guiana: 4 mi. east of Georgetown
(H. G. Kugler). Durca Guiana: Corentyn River (H. G. Kugler).

Melongena patula Broderip and Sowerby
Plate 99

Pyrula melongena fusco-nigricans Sowerby 1825, Catalogue of the Shells in the Collection of the Earl of
Tankerville, London, p. 62 [nomen nudum].

Qcr

Pyrula patula Broderip and Sowerby 1829, Zoological Journal 4, p. 377 (Mazatlan, Mexico).

Eastern Pacific JOHNSONIA, No. 35 169

Cyrtulus patula Broderip and Sowerby, Jay 1850, Catalogue of the Shells contained in the collection of John
C. Jay, New York, 4th edition, p. 323.

Cassidulus patula Broderip and Sowerby, Mérch 1852, Catalogus Conchyliorum Comes de Yoldi, Denmark,
p. 103.

Melongena patula Broderip and Sowerby, Petit de la Saussaye 1852, Journal de Conchyliologie 3, p. 146.

Galeodes patula Broderip and Sowerby, Zetek 1918, Revista Nueva, Panama 5, p. 526 [p. 21 in reprint].

Galeodea patula Broderip and Sowerby, Pilsbry and Lowe 1932, Proceedings Academy Natural Sciences,
Philadelphia 84, p. 114.

Description. Shell reaching 260 mm. (about 10 inches) in length and 190 mm. (about
74 inches) in width, solid and usually smooth or only moderately spinose on the whorl
shoulder. Color generally of a dark chestnut-brown with a light cream-colored band just
below the widest bulge of the whorl. This band may consist of several very narrow bands
grouped together. In addition, there may be other spiral bands of the same color, usually
thread-like and varying greatly in number. A single specimen that we have seen also
shows axial streaks of this cream color. Whorls 74 to 8, strongly convex, particularly
above the periphery. Spire somewhat elevated in young specimens, but in the adult stage
the spire is depressed, occasionally hardly extending above the top of the body whorl.
Aperture subelliptical with a rather deep anal notch and a broad, flattened and short
siphonal canal. Parietal wall rather heavily glazed. Palatal or outer lip moderately thin
and only moderately crenulated. Columella nearly perpendicular and twisted. Umbilicus
closed. Suture deeply channeled; occasionally, the later whorls are folded over the early
whorls. Sculpture consisting of a single row of spines at the whorl shoulder or none at

Plate 99. Melongena patula Broderip and Sowerby. Figs. 1-3. Guaymas, Sonora, Mexico. Figs. 1 and 3, about
4/5x. Fig. 2. Young specimen (about 1.3x).

170 JOHNSONIA, No. 35 Melongena

all. Most specimens show numerous and rather fine, spiral, incised grooves developed
mainly on the lower half of the shell. Young specimens and the early whorls of adult
specimens are usually strongly nodulose. Operculum horny, unguiculate, rather solid
with the nucleus at the lower end and sculptured with rather strong concentric lines of
growth. The inner surface has a large muscle scar which has a smooth shiny margin and
2a somewhat roughened and dull central area. Periostracum quite smooth, dull and hav-
ing exceedingly fine axial growth lines.

length width
260 mm.* 190 mm. Panama
195 “a 144 Panama
131 - 98 Mazatlan, Mexico

* early whorls lost

Types. ‘Type specimens of this species are probably in the British Museum. The type
locality is Mazatlan, Mexico.

Remarks. See also Remarks under Melongena melongena Linné.

It is possible that this species was derived from the West Indian Melongena and en-
tered the Pacific from the Atlantic during the submergence of the Isthmus of Panama
in the late Miocene and early Pliocene times. Today the two forms are differentiated
sufficiently to be considered separate species. There are many similar cases of such paral-
lel species in the two oceans, such as Strombus gracilior Gray, Purpura patula panza
Gould, Barnea subtruncata Sowerby, and Pholas chiloensis Molina of the Pacific and
Strombus pugilis Linné, Purpura patula Linné, Barnea truncata Say and Pholas campe-
chiensis Gmelin of the Atlantic.

Range. Gulf of California from Guaymas, Mexico south to Bahia de Caraques, Ecua-
dor. ‘This southern record is given by Bayer (1952, p. 272).

Specimens examined. MExico: Guaymas, Sonora (H. R.'Turver; A.Sorensen); Maz-
atlan(MCZ). Nicaracua: Corinto (Charleston Museum). Cosra Rica: Gulf of Nicoya
(MCZ). Panama: Chame Bay; Canal Zone (both MCZ); Verrado Beach (C. M. Dum-
bauld); near Balboa (J. Zetek; MCZ).

Subgenus Rexmela Olsson and Harbison

Revmela Olsson and Harbison 1953, Academy of Natural Sciences Philadelphia, Monograph 8, p. 213.

Type species, Melongena subcoronata Heilprin (= Melongena corona Gmelin), original
designation.

Shell moderate in size to fairly large, generally solid and strong but not massive. Spire
extended, the later whorls not enveloping the earlier whorls. Suture not channeled but
coarsely imbricate, the imbrications being previous margins of the anal canal. Spines vari-
able, claw-like to spike-like, generally open on their forward margin. ‘They may be pro-
duced at right angles to the shell surface but generally they are somewhat parallel to the
long axis of the shell, and frequently curve inwardly.

Rexvmela was introduced as a subgenus, separating the Pliocene Melongena subcoronata
Heilprin from the recent M. corona Gmelin. In our opinion, these two forms are syn-
onymous; in fact, we can duplicate almost exactly the type of MW. subcoronata with recent
specimens from Lossman’s Key.

Western Atlantic JOHNSONIA, No. 35 171

However, the name Rexmela is available to use for a subgenus separating corona, its
subspecies and closely related forms, from M. melongena Linné and M. patula Broderip
and Sowerby, two species quite different from the corona complex.

This subgenus is limited in its distribution to Florida, Alabama and the northern tip
of Yucatan.

The Melongena corona complex

The multiplicity of forms in Melongena corona Gmelin is quite probably due to factors
of isolation of smaller or larger areas along the central and southwest coast of Florida
during its geologic history since the Pliocene. This is a region of very low relief and even
minor changes in the oceanic level would affect profoundly much of the coast line. Areas
of land were invaded by the sea as the oceanic level was raised, new habitats were created
and colonies of this species developed as they moved into these new areas. Possibly
chance alone brought in a few individuals with a similar gene complex and thus a popu-
lation evolved that was remarkably uniform. In others, one or more different assemblages
invaded a new area and a mixture took place giving rise to a very varied population. A
lowering of the oceanic level would have a similar effect, exposing a new land area that
would shortly build up the mangrove association or a similar assemblage of a somewhat
brackish water fauna and flora.

Plate 100. Melongena corona corona Gmelin. Figs. 1-3. Barfield Bay, Caxambas, Florida. Figs. 4-5. Pinaire,
Pine Island, Lee Co., Florida. Fig. 6. Goodland, Marco, Florida (all 5/6x).

172 JOHNSONIA, No. 35 Melongena

Naturally such changes were slow and as they took place, either as a plus or minus
factor of oceanic level, the fauna and flora advanced or retreated. Thus the present situ-
ation is perhaps but a picture of much of the past. We are but viewing at this time a
medley of colonies or populations that have been forced to move back and forth; some
maintaining a certain genetic purity, others becoming thoroughly mixed by the vicissi-
tudes of chance introductions.

The area of greatest disturbance appears to be between Tampa Bay and Cape Sable.
North of Cedar Keys Me/ongena exhibits a fine example of a cline, ending at Gulf Shores,
Alabama as a subspecies which we have called johnstonei. Starting at Cape Sable and
going east and then north along the east coast of Florida the populations of Melongena
are very uniform and are considered here as the subspecies a/tispira. The Keys, from Key
Largo to the Tortugas possess another species, Melongena bicolor Say, which is remark-
ably uniform.

Melongena corona corona Gmelin
Plates 94-97: 100-103

Murex corona Gmelin 1791, Systema Naturae, ed. 13, p. 3552 (Sinu Mexicano).

Melongena belknapi Petit de la Saussaye 1852, Journal de Conchyliologie 3, p. 65, pl. 2, fig. 5 (coast of
Florida).

Melongena subcoronata Heilprin 1887, Transactions Wagner Free Institute of Science, Philadelphia 1, p. 70,
pl. 1, fig. 3 (Pliocene: Banks of the Caloosahatchie River below Fort Thompson, Florida).

Melongena corona aspinosa Dall 1890, Transactions Wagner Free Institute of Science, Philadelphia 3, p. 120,
pl. 9, fig. 6 (Pliocene: Caloosahatchie Beds, Florida).

Melongena corona inspinata Richards 1933, Nautilus 47, p. 57, pl. 6, fig. 1 (near Sarasota, Florida).

Melongena perspinosa Pilsbry and Vanatta 1934, Nautilus 47, p. 120, pl. 12, fig. 1 (Lossman’s Key, near
Shark River, Florida).

Melongena corona incurvata Lermond 1936, Check List of Florida Marine Shells, Gulfport, Florida, p. 40
[nomen nudu m).

Melongena corona sargenti Emery and Lermond 1936, Check List of Florida Marine Shells, Gulfport, Florida,

p. 42 [nomen nudum |.

Plate 101. Melongena corona corona Gmelin. Fig. 1. Paratype, Melongena corona inspinata Richards ( = Melon-
gena corona corona Gmelin) Palma Sola Key, Florida (natural size). Fig. 2. Boca Ciega Bay, Gulfport,
Florida (about 1.3x).

Western Atlantic JOHNSONIA, No. 35 173

Melongena corona trinodulosa Emery and Lermond 1936, Check List of Florida Marine Shells, Gulfport,
Florida, p. 40 [nomen nudum].

Melongena corona mulletensis ‘Lermond’ VanHyning 1940, A Check list of the Mollusca of Florida, Gaines-
ville, Florida, p. 19 [nomen nudum).

Melongena corona perspectiva Abbott 1954, American Sea Shells, New York, p. 72. [This was a lapsus for
Melongena corona perspinosa Pilsbry and Vanatta. |

Description. Shell reaching 205 mm. (about 8 inches) in length, fairly thin to relatively
thick and solid, and usually sculptured with spines. Color generally ivory with spiral
bands of light orange-brown to dark red-brown. These bands are very variable, some
specimens having almost a continuous series of bands of varying widths. However, the
most widespread pattern consists of three major bands: one at the whorl shoulder, one
about at the whorl periphery and a broad band at the base of the whorl. Whorls 7, rather
convex and shouldered. The shoulder is usually horizontal and rather narrow: occasion-
ally it may be broad and with a downward slope. Spire subdepressed to extended. A per-
ture subovate. Outer lip thin and usually crenulated. Inner lip usually consisting of a
thin callus, but in old specimens it becomes thick and white in color. Columella rather
broad and twisted. Umbilicus glazed over by the parietal shield. Suture relatively incon-
spicuous, generally a little more pronounced on the earlier whorls. Sculpture exceedingly
variable. Usually there is a single row of rather large spines at the whorl shoulder and
another at the base of the whorl. However, occasional colonies may have specimens ex-

Plate 102. Melongena corona corona Gmelin. Holotype, Melongena perspinosa Pilsbry and Vanatta (= M. corona
Gmelin), Lossman’s Key, Florida (about 1. 1x).

174 JOHNSONIA, No. 35 Melongena

hibiting the large shoulder spines and one to two rows of smaller spines immediately
below: in addition, there is sometimes a row of smaller spines between the shoulder series
and the suture. In other colonies the spines may be reduced to a single row at the whorl
shoulder, the basal row being absent. In various colonies the spines may project at right
angles to the shell axis, or more commonly project upwardly and may even be arched
toward the spire. There is a great deal of variation in the number of large shoulder spines,
these ranging from 10 to 26 on the body whorl. Generally the spines are claw-like in
appearance and are open on the forward side. Occasionally these spines are closed, at
least near the base, especially in colonies from the region between Cape Romano and
Lossman’s River. At the suture there is a series of imbrications representing the pre-
vious growth stages of the anal canal. These are most conspicuous on the last three
whorls. In addition to the spiral rows of spines there are numerous spiral ridges which
vary in their intensity between colonies. The early whorls are generally rather strongly
axially costate. Operculum unguiculate, the nucleus basal, the outer surface with nu-
merous and exceedingly fine concentric ridges. Inner surface with the palatal margin
smooth and glazed, and the inner area with a series of rather widely separated concentric
ridges which appear shingled. Periostracum usually colored a dark and dull greenish-
brown to a dull red-brown.

length width *
205 mm. 127mm. Goose Cove, Cedar Keys, Florida
187 123 Big Carlos Pass, Sanibel Island, Florida
138 95 Lossman’s Key, Florida
98 70 Holotype of M. perspinosa P. and V.

* not including spines

Types. It is practically certain that Gmelin had no specimens of this species, his de-
scription being based on the Latin diagnosis and figures in Chemnitz. We here select
Chemnitz’s figure in the Conchylien-Cabinet 1788, (1) 10, pl. 161, fig. 1527 as the type
figure. The type locality was given originally as the Gulf of Mexico. We here restrict
it to Tampa Bay, Florida, as possibly the area from which this material may have come.
The type of Melongena belknapi Petit de la Saussaye is in the collection of the Journal
de Conchyliologie. The holotypes of MW. subcoronata Heilprin and M. perspinosa Pilsbry
and Vanatta are in the Academy of Natural Sciences, Philadelphia. The holotype of WZ.
corona inspinata Richards is in the collection of Mrs. E. R. Edgerly, Trenton, New Jer-
sey; paratypes are in the Academy of Natural Sciences, Philadelphia. The holotype of
Melongena corona aspinosa Dall is in the United States National Museum, no. 135319.

Remarks. The lite history of this species is exceedingly interesting and much of it ac-
counts for the variation exhibited by the typical form. Through the kindness of Dr. and
Mrs. David Schmidt we were able to get live material for our studies. Melongena corona
breeds in the summer months, starting in early July. Live specimens were received in
March and placed in an aquarium. They continued to be active throughout the spring
and on July 11, laid the first egg capsules. Their gross embryology was followed daily.
Each day a capsule was cut open and the young examined. In this way a study was made
from the initial egg stage to the young snail that emerged from the capsule, as described
in the introduction. All stages in the development are passed within the capsule, from
the egg through the trochophore and veliger larvae, and the young finally emerge as very

Western Atlantic JOHNSONIA, No. 35 17

Gr

small but nevertheless fully developed crawling snails. Naturally, egg capsules attached
to objects subject to mechanical movement may be dispersed, but in the main, the young
are very closely associated with the region where they were produced. From a purely
genetic standpoint this would account for the general uniformity of many colonies and
also for the variation shown by other colonies. Species which possess free-swimming lar-
vae are generally quite constant in their characters over most of their range, much de-
pending, of course, on the length of larval life and the ability of water currents as a dis-
persal factor.

According to Dall (1890, p. 118) various members of the Melongenidae appeared in
Americain the Upper Eocene. So far as we can trace Melongena appeared in the Pliocene
of central Florida. ‘There seems to be no difference between the recent Melongena corona
corona Gmelin and the Pliocene M. subcoronata Heilprin. Our figure of aspecimen from
Lossman’s Key, Florida (Plate 103, fig. 1) is almost an exact replica of the figure given
by Heilprin for his swbcoronata. We have other specimens from Fort Myers Beach,
Florida which are also nearly identical with Heilprin’s figure.

Range. The subspecies M. c. corona extends from Keaton’s Beach, Taylor County,
Florida south to the north end of Cape Sable, Florida.

Specimens examined. FLortpA: Lossman’s Key, Ten Thousand Islands (D. and N.
Schmidt; MCZ; USNM): Plover Key, 5 mi. north-northwest of Lossman’s River,
Monroe Co.; Allen River, Everglades, Collier Co. (both MCZ); Pavilion Key (Carne-

Plate 103. Melongena corona corona Gmelin. Fig. 1. Lossman’s Key, Ten Thousand Islands, Monroe County,
Florida (1.1x). Fig. 2. Holotype, Melongena subcoronata Heilprin (= M. corona Gmelin) from Heilprin 1887,
pl. 1, fig. 3 (about natural size).

176 JOHNSONIA, No. 35 Melongena

gie Museum): Cape Romano (D. and N. Schmidt): Horse Key, 14 miles southeast of
Marco, Collier Co. (MCZ): Marco Island (D. and N. Schmidt: R. Merrill: J.Schwengel:
M. Branham): Gordon River, Naples (D. and N. Schmidt): Barfield Bay, Naples (R.
Cahoon: D.and N. Schmidt): Bonita Beach, Bonita Springs (MCZ: D. and N. Schmidt):
Estero Island (R. Merrill: D. and N. Schmidt: USNM): Punta Rassa (R. Merrill: D.
and N. Schmidt: USNM; MCZ): Caloosahatchee River, Port Comfort (D. and N.
Schmidt): Mouth of Alligator Creek, Charlotte Harbor (MCZ): Sanibel Island (J.
Schwengel: R. Merrill: USNM: MCZ): Causeway to Mattacha Pass and Pine Island
(G. Weber): Bokeelia Beach, Pine Island (D. and N. Schmidt: G. Weber): Pineaire,
Pine Island: St. James, Pine Island (both D. and N. Schmidt): Boca Grande (H.
Dodge): Gasparilla (Florida State Museum): Johnson Cove and Cedar Point, Lemon
Bay (both MCZ): Englewood Beach (R. Merrill): Venice: Casey's Key. Manatee Co.
(both Carnegie Museum): Mouth of North Creek, Little Sarasota Bay: Upper end of
Little Sarasota Bay, 54 miles south of Sarasota (both MCZ): Sarasota (MCZ: R. Ca-
hoon): Manatee River (MCZ): Palma Sola Bay, Manatee Co.: Anna Maria Key,
Bradenton (both D. and N. Schmidt): Long Key, near Pass-a-grille: Mullet Key. St.
Petersburg: Clam Bayou, Boca Ciega Bay, Gulfport: Big Bayou and Monte Cristo Is-
land, St. Petersburg: Piney Point, Lower Tampa Bay (all MCZ): Rocky Point and
Satety Harbor, ‘Tampa Bay (both Florida State Museum): Seminole Pier. Clearwater
Bay (G. Weber): Davis Causeway, Clearwater (MCZ): St. Josephs Sound, Dunedin
(G. Weber): Tarpon Springs (J. Weber): Bayou between Green Key and Port Richey
(G. Weber): Crystal River (H. and K. Johnstone): Sea Horse Key, Cedar Keys (MCZ):
Goose Cove, Cedar Keys (S. Whitman): Horseshoe Point, Dixie Co. (T. Pulley):
Keatons Beach (D. and N. Schmidt).

Melongena corona altispira Pilshry and Vanatta
Plate 104

Melongena corona altispira Pilsbry and Vanatta 1934, Nautilus 47, p. 119 (Oceanus, Brevard Co., Florida).

Description. Shell reaching 72 mm. (about 3 inches) in length, thin but strong, and
generally sculptured with spines. Color generally ivory with chocolate brown bands, one
at the whorl shoulder, the second at the whorl periphery, and the third at the base of the
whorl. These bands appear to be quite constant. Whorls 7, convex and shouldered. The
shoulder is usually horizontal and narrow. Spire somewhat elevated. Aperture subovate.
Outer lip thin and finely crenulate. Inner lip consisting of a rather thin callus. Columella
broad and twisted. Umbilicus glazed over by the parietal shield. Suture relatively incon-
spicuous. Sculpture variable. In this form there is usually but a single row of small and
rather numerous spines at the whorl shoulder. However, many specimens occur, particu-
larly in the Cape Sable area, that are devoid of spines, though others in the same colony
may have a single row at the whorl shoulder. ‘These spines project upward and are sel-
dom arched. An occasional specimen will show a row of weak and small spines at the
base of the shell. ‘There are numerous and fine spiral ridges which are a little stronger near
the base. The early whorls are strongly and axially costate. Sutural imbrications are
only weakly developed. Operculum similar to that of the typical form. Periostracum
very thin and a dull brownish in appearance.

Western Atlantic JOHNSONIA, No. 35 177

length height whorls

71.0 mm. 41.0 mm. 8 Mosquito Lagoon, Oak Hill, Florida
(025 44.0 il Matanzas Inlet, St. Augustine, Florida
70.0 40.5 8 Cocoa, Florida

58.0 33.5 7 7 miles W. of Flamingo, Florida

Types. The holotype of Melongena corona altispira Pilsbry and Vanatta is in the
Academy of Natural Sciences, Philadelphia, no. 72490. The type locality is Oceanus,
Brevard County, Florida.

Remarks. This subspecies does not appear to be abundant anywhere throughout its
range to judge from the collections we have seen. It appears to be a direct offshoot of
M. corona corona. Actually, the type series described by Pilsbry and Vanatta are atyp-
ical for the subspecies. They appear to be specimens that may have existed under hyper-
saline conditions, a habitat which frequently causes distortion in shell structure.

We possess other specimens of this subspecies from Oceanus that are very similar to
the normal form which exists along the entire east coast of Florida. They were obviously
not from the same ecological spot.

Many specimens of this subspecies are devoid of spines, others are weakly spinose and
in still others the shoulder spines are exceedingly irregular in their development. The
basal row of spines is weak and generally lacking.

We have seen specimens at two localities on the southwest coast that apparently be-
long to this subspecies. These reached Marco Island and Cape Romano probably by
drifting from the Cape Sable region.

This subspecies differs from typical corona by being on the average a little smaller,
having but a single row of shoulder spines which are weakly developed and frequently
absent. The basal row of spines on altispira is usually lacking, but when it is present
there are always shoulder spines in addition.

Plate 104. Melongena corona altispira Pilsbry and Vanatta. Fig. 1. Holotype, Oceanus, Florida. Fig. 2. Ba-
nana River, 2 miles east of Coco Beach, Florida. Fig. 3. New Smyrna, Florida (all about 8x).

178 JOHNSONIA, No. 35 Melongena

We have not detected any evidence of a true cline along the east coast of Florida in
this subspecies. Perhaps the data at hand are still insufficient. On the other hand, until
the inland water way was made continuous, there were many stretches, some long ones,
that offered only the open ocean as a highway, areas quite unsuitable for Melongena and,
in addition, having coastal currents that trend southwards rather than to the north. Our
most northern record, Matanzas Inlet, was based upon a single dead shell and this may
well be a recent introduction into this area. The Florida portion of the inland waterway
needs far more study, both in the dredged cuts as well as along the entire Indian **River”’
area where only a boat channel was dredged.

See also Remarks under ©. bicolor Say.

Range. This subspecies extends from the region of Cape Sable, Florida in the south,
to the east and north as far as Matanzas Inlet near St. Augustine.

Specimens examined. FLORIDA: Matanzas Inlet, 15 miles south of St. Augustine;
Daytona Beach; New Smyrna (all MCZ):; Mosquito Lagoon, near Oak Hill (USNM);
Indian River, Cocoa, Brevard Co. (USNM; MCZ; J. Schwengel):; Ocean Beach, Bre-
vard Co. (Carnegie Mus.): Oceanus, Brevard Co. (ANSP: MCZ); Eau Gallie(MCZ);
Sebastian (R. Merrill); St. Lucy Inlet (J. M. Cannon); Virginia Key, Miami (J. Weber);
Matheson Hammock, Biscayne Bay (R. Merrill): Coral Gables (J. Weber; R. Merrill);
Coconut Grove: Key Largo Station, Key Largo (both MCZ); Barnes Sound, Key Largo
(J. Weber); Madeira Bay (MCZ): 7 miles west of Flamingo, Cape Sable (D. and N.
Schmidt): Sandy Key, Cape Sable(MCZ; J.Schwengel): Marco Island; Cape Romano
(both D. and N. Schmidt).

Melongena corona johnstonei, new subspecies
Plates 96 and 105

Description. Shell reaching 171 mm. (6# inches) in length, rather solid and strongly
sculptured. Color generally pale ivory overlaid with two or three bands of dark chocolate-
brown which coalesce in the adult forming an almost solid brown shell. In some speci-
mens the brown bands persist in the adult and these are best observed in transmitted
light as seen from within the aperture. Whorls 74 to 8, rather strongly convex and
shouldered. Spire extended. Aperture subovate. Outer lip fairly thin and rather finely
crenulate. Inner lip usually consisting of a thin to fairly thick callus. Columella broad
and twisted. Umbilicus glazed over by the parietal shield. Suture relatively inconspicu-
ous. Sculpture consisting usually of a single row of strong, erect or recurved spines on
the margin of the whorl shoulder. Rarely there exists a rather weak series of spines near
the base of the shell. In addition to the spines there are numerous fine spiral ridges which
are strongest near the base. At the suture there are numerous imbrications representing
previous growth stages of the anal canal. Operculum as in the typical form. Periostra-
cum thin and a dull greenish-gray in color.

length width whorls
171 mm. 94.0 mm. 8.0 Paratype
126 76.0 8.0 Port St. Joe, Florida
102 54.0 to Holotype
94 56.5 (GE Pensacola, Florida
83 51.0 7.0 Panacea, Wakulla Co., Florida

Western Atlantic JOHNSONIA, No. 35 179

Types. Holotype, Museum of Comparative Zoélogy, no. 189687, from Little Lagoon,
Gulf Shores, Alabama. Paratypes from the same locality in the Museum of Compara-
tive Zodlogy, the United States National Museum, the Academy of Natural Sciences,
Philadelphia, the Florida State Museum, the Alabama Museum, and the Collections of
Harry and Kathleen Johnstone, David and Nevada Schmidt, and D. Thaanum.

Remarks. ‘This subspecies has a very limited distribution in northern Florida and ex-
treme eastern Alabama. It appears to merge more or less with the typical form in the
region of Apalachee Bay, Florida. It differs in being more elongate, in being in general
much darker in color, and in having a brown coloration well diffused over most of the
shell leaving a few narrow, spiral bands of white. The shoulder is generally broader than
in M. c. corona, and the shoulder spines are vertical and more generally pointed inwardly
toward the spire.

At Little Lagoon, Gulf Shores, Alabama, M. c. johnstonei were feeding on Ensis
minor Dall and Tagelus divisus Spengler. They would locate these bivalves and then
extend their proboscises down into the burrow and clean them out. Dead specimens of
these two species were the only common bivalves in the beach drift and Melongena corona
Johnstonei was the only large gastropod that we collected along two miles of shore line.

This subspecies exhibits a smooth cline. The gradient of change starts above or along
the area of Dead Man’s Bay, which is the ‘‘bight’’ area of north central Florida on the
west coast. Transitional elements with M. c. corona appear in this region. As colonies

Plate 105. Melongena corona johnstonei Clench and Turner. Fig. 1. Holotype. Figs. 2-3. Paratypes, all Little
Lagoon, Gulf Shores, Alabama (all natural size).

180 JOHNSONIA, No. 35 Melongena

are obtained at stations both north and west, the specimens become somewhat larger and
definitely more attenuate, reaching their ultimate size and attenuation at Gulf Shores,
Alabama. The scant data available indicate that if WZ. corona occupied this territory dur-
ing the warm Pliocene, it was killed out during the cold Pleistocene and invaded this
territory anew after the Pleistocene. The subspecies johnstone:, having a more uniform
gene complex has become somewhat stabilized in the area north of Cedar Keys where it
is more or less isolated from the typical M. c. corona. This is a result of the small tidal
flow in the vicinity of Cedar Keys so that there has been far less opportunity for variable
genetic elements of the corona complex of the south to reach this area by mechanical
means.

See also the remarks under both M. c. altispira and M. bispinosa.

It is with pleasure that we name this subspecies for Mr. Harry I. Johnstone of Mobile,
Alabama.

Range. From Gulf Shores, Alabama east to Panacea, Wakulla County, Florida.

Specimens examined. ALABAMA: Little Lagoon, Gulf Shores (Mr. and Mrs. H. I.
Johnstone: MCZ). Froripa: Pensacola (MCZ; T. Pulley); west end of Santa Rosa Is-
land (Univ. of Florida; J. Weber); Fort Walton, Santa Rosa Island (H.1. Johnstone):
Port St. Joe (R. Merrill; D. and N. Schmidt: Univ. of Florida); James Island, Frank-
lin Co. (T. Pulley); Panacea (MCZ). |

Melongena bispinosa Philippi
Plate 106

Pyrula bispinosa Philippi 1844, Abbildungen und Beschreibungen Conchylien 1, Pyrula, p. 2, pl. 1, figs.
7-8 (locality unknown).

Pyrula martiniana ‘Pfeiffer’ Philippi 1844, Abbildungen und Beschreibungen Conchylien 1, Pyrula, p. 2,
pl. 1, fig. 9 (locality unknown).

Melongena bispinosa Philippi, Petit de la Saussaye 1852, Journal de Conchyliologie 3, p. 157, pl. 8, fig. 3.

Melongena corona bispinosa Philippi, Pilsbry 1934, Nautilus 47, p. 120.

Description. Shell reaching 75 mm. (about 3 inches) in length, rather light in struc-
ture and sculptured. Color light brownish-yellow with one to three spiral bands of dark
brown; interior of aperture light yellow. Whorls 6 to 7 and shouldered. Spire somewhat
extended ; aperture subovate; outer lip rather thin and crenulated. Inner lip usually con-
sisting of a fairly thick callus on the parietal wall. Columella rather broad and flatly sig-
moid in profile. Umbilicus usually closed, occasionally with a small perforation. Suture
well defined. Sculpture consisting usually of one to three rows of spines at the whorl
shoulder; in addition, there is a single row of scale-like spines midway between the peri-
phery and the base of the shell. At the base there are a series of scales which represent
former margins of the siphonal canal. A similar series of small scales exists at the suture,
which are previous margins of the anal canal. There are numerous thread-like spiral]
ridges. Axial costae rather strongly developed on the early whorls. These costae are
somewhat nodulose.

length width
75 mm. 50.0mm. from Petit de la Saussaye
43, (De

26.5 Dzilam, Yucatan, Mexico
42 24.5

6 6é 6e

Western Atlantic JOHNSONIA, No. 35 181

Types. 'The whereabouts of the types of both Melongena bispinosa and M. martiniana
Philippi is unknown. They are not in the British Museum (Natural History), where
some of Philippi’s shells are located, according to Mr. Guy Wilkins who kindly checked
this for us.

Remarks. This species appears to be nearest in relationship to certain forms of M. c.
altispira of the Florida east coast. It differs from all other members of the corona com-
plex by possessing scale-like spines behind the siphonal canal. These are actually previous
margins of this canal. When the animal enlarges the shell it builds a completely new
margin to the canal and thus the older ones that are left behind project outwardly, giv-
ing the appearance of scales. ‘This same morphological structure is of frequent occurrence
in Murex and Rapana.

The occurrence of this species in Yucatan is one of those anomalous problems in dis-
tribution that are exceedingly difficult to explain. This species is completely isolated from
other members of the corona complex and even in Yucatan occupies only a small area so
far as is now known.

It was originally described without any known locality and later Petit de la Saussaye
described and figured a specimen which he received from Largilliert, also without local-
ity. Largilliert had collected rather extensively in Central America, including Yucatan,
about 1842.

In 1890, Angelo Heilprin, a member of the staff of the Academy of Natural Sciences
in Philadelphia, made an expedition to Yucatan mainly in quest of fossil material. Re-
cent marine shells were also collected and these were published by F. C. Baker (1891).
Heilprin collected M. bispinosa at two stations, Progresso and Silam [Dzilam de Bravo],
both on the northern coast of the peninsula. We have seen an additional lot labeled only
Yucatan. Undoubtedly more collecting may extend the known range of this species.

Perhaps M. bispinosa is only a remnant from the Pliocene of a once far more widely
distributed element of the corona complex. During this warm period in the geologic past
this species complex may have occupied the entire Gulf coast area and later, during the
cold Pleistocene these forms may have been driven south or were killed out in the north-
ern portion of the Gulf region. By this means, M. bispinosa remains as a relict species,
completely isolated from other elements to which it is obviously closely related.

See also remarks under M. corona johnstonei.

2

Plate 106. Melongena bispinosa Philippi. Figs. 1-2. Yucatan (natural size).

182 JOHNSONIA, No. 35 Melongena

Range. Coast of Yucatan from Campeche (N. E. Weisbord, 1926) to Dzilam de Bravo.

Specimens examined. Mexico: Dzilam de Bravo and Progresso, Yucatan (both ANSP).

Melongena bicolor Say
Plates 96; 107-108
Fusus bicolor Say 1827, Journal Academy Natural Sciences, Philadelphia 5, p. 215 (southern coast of East
Florida).
Hemifusus corona minor Sowerby 1878 [1879], Proc. Zoological Society London, pp. 796, 798 (plate caption),
plate 48, fig. 13 (Key West, Florida).
Hemifusus corona estephomenos Melville 1881, Journal of Conchology 3, p. 157 (Key West, Florida).

Description. Shell reaching 50 mm. (about 2 inches) in length, thin but strong and
sculptured with spines, Color a light ivory to pinkish-ivory with spiral bands of brown.
These bands are variable but generally there are three, one at the whorl! shoulder, one at
the periphery of the whorl and a third at the base of the whorl. Early whorls often a
uniform dark brown. There are occasional specimens without bands. Whorls 8, moder-
ately convex and with anarrow shoulder. Spire elevated. Aperture subovate. Outer lip
thin and very finely crenulated. Inner lip consisting of a very thin glaze. Columella
broad and twisted. Umbilicus glazed over by the parietal shield. Sculpture somewhat
variable. On the whorl shoulder there may be a single row or rarely two rows of rather
small and close-set spines which are erect. Very rarely a basal row of spines is present.
In addition to the spines, there are numerous very fine spiral ridges which are a little
stronger near the base of the whorl. Axial costae strong and well developed on the early
whorls, sometimes persisting even on the body whorl. Sutural imbrications very small
and only weakly developed. Operculum as in M. corona. Periostracum very thin and a
dull brownish in color.

length height whorls

49.0 mm, 25.0 mm. 8 Key Vaca, Florida
43.5 23.0 f Salen ss vf
43.0 OO 5 8 Tortugas, Florida

Types. The holotype of Melongena bicolor Say is in the Academy of Natural Sciences,
Philadelphia, no. 84276, from the southern coast of east Florida, Mr. Elliott collector.
The types of M.c. minor Sowerby and M.c. estephomenos Melvill, both names based upon
the same specimens, are probably in the British Museum. The type locality is Key West,
Florida. This last can also be accepted as the type locality for bicolor Say.

Remarks. This is quite a distinct species. In relationship it appears to be nearest to
M. c. altispira Pilsbry and Vanatta but differs by being smaller, much lighter in color
and having more and _ better developed shoulder spines. We have seen no hybrid speci-
mens even where their ranges overlap or approximate one another in the vicinity of Bis-
cayne Bay. In addition, the ecology of the two forms is quite different, M. bicolor living
usually on the outer sandy beaches where completely marine conditions exist, while ©.
c. altispira is found in brackish water areas. Young specimens of the two species may be
difficult to differentiate.

This species is remarkably uniform in size, specimens from several localities showing
but little variation. The number of spines on the body whorl, though variable, appears

Western Atlantic JOHNSONIA, No. 35 183

to be far more constant than is the case in M. corona. There is no question but that Say’s
name bicolor applies to this species. His description was based upon very young and worn
specimens, but none the less, it agrees entirely with the young of the form that occurs
on the Lower Florida Keys (Plate 108, figs. 1-2). Say’s type locality ‘“‘Southern coast
of East Florida’ could certainly apply to the Florida Keys, and may possibly have been
Key West.

Our northernmost record is Biscayne Bay and from here the records extend down the
Keys generally on the outer sandy beaches to as far as the Dry Tortugas.

Mr. John H. Butler has kindly contributed the following data on certain of their habits.
“These Melongena bicolor are to be found only on the Atlantic side of the Florida Keys.
They have developed a method of protection from the elements which shields them from
hot sun or wave action during storms. They bury themselves in the marly sand and take
their food along with them. As the tide comes in they emerge when covered with a few
inches of water, then they hunt their meal. They gather two or three Cerithium, carry
these around until they decide it’s time to dig down again, then they go completely out
of sight, sometimes as much as four inches down. There they leisurely devour their lunch
and wait for the tide to come back again. They do not seem to like brackish water, for
they will not come out ona rainy day. Caught out in a summer shower they at once dig
down, and most of them will stay down until the tide turns again.

‘*T have found no live specimens on the Gulf side of the Keys; a few dead shells occa-
sionally, more often than not inhabited by hermit crabs. Scattered specimens may be
found in some of the harbors and back bays of the Atlantic side, such as Boot Key Har-
bor off Key Vaca. These have most likely been washed in by a storm for they do not
thrive in such locations. They seem to prefer the open Atlantic shores where the sand is

Plate 107. Melongena bicolor Say. Figs. 1-4. Outer beach, Grassy Key, Lower Keys, Florida. Figs. 5-7.
Lower Matecumbe Key, Lower Keys, Florida (all 1.2x).

184 JOHNSONIA, No. 35 Melongena

heavy with a high marl content. They are entirely absent from such Keys as Bahia Honda
having a fine clean sand beach. Such sand moves readily in a storm and they would be
washed up on the beach and exterminated. They seem to be strictly carnivorous, prefer-
ring their food fresh and alive. I have never found them feeding on dead fish or dead
crabs, unlike the large Melongena corona of the west Florida Coast. They are absent
from large areas where there are no Cerithium, other conditions apparently favorable.
Where Cerithium are scarce you will find adults only, no juveniles. ”’

Range. From Biscayne Bay southwest along the Keys to Dry ‘Tortugas.

Specimens examined. FLoripa: near Coral Gables Canal, Biscayne Bay (D. Moore);
Plantation Key (D. and N. Schmidt; H. Moore); Windleys Key (D. and N. Schmidt);
Upper Matecumbe Key (MCZ); Lower Matecumbe Key (Carnegie Mus.; MCZ; J.
Weber): Upper Grassy Key (D. and N. Schmidt); Grassy Key (J. Butler; R. Cahoon;
I). and N. Schmidt); Crawl Key (D. and N. Schmidt); Key Vaca (J. Butler; D. and
N. Schmidt); Bahia Honda Key (R. Merrill); Torch Key (MCZ): Sugar Loaf Key
(MCZ; Carnegie Mus.: D.and N. Schmidt); Boca Chica Key (Charleston Mus. ): Stock
Island (Carnegie Mus.):; Key West: Dry Tortugas (both MCZ).

Plate 108. Melongena bicolor Say. Fig. 1. Lectotype, Fusus bicolor Say (= Melongena bicolor Say), southern
Florida. Fig. 2. Key Vaca, Florida (young) (both about 5.2x).

Genus Pugilina Schumacher
Pugilina Schumacher 1817, Essai d’un Nouveau Systéme des Habitations des Vers Testacés, py 216.
Type species, Pugilina fasciata Schumacher (= Murex morio Linné), monotypic.

Shell subdepressed to attenuate, generally rather heavy in structure and relatively
smooth to strongly nodulose. Periostracum dark brown and relatively thick. Spiral
sculpture variable, consisting of numerous threads or incised lines.

Western Atlantic JOHNSONIA, No. 35 185

Subgenus Pugilina Schumacher

Shell subdepressed to extended, rather solidly constructed, smooth to strongly nodu-
lose and having relatively weak spiral threads.

Pugilina morio Linné
Plates 96 and 109

Murex morio Linné 1758, Systema Naturae, ed. 10, p. 753 (locality unknown); ibid. ed. 12, 1767, p. 1221
(M. Africano); Hanley 1855, Ipsa Linnaei Conchylia, p. 300.

Fusus morio Linné, Réding 1798, Museum Boltenianum, p. 120.

Fusus coronatus Lamarck 1803, Annales du Muséum d’Histoire Naturelle, Paris 2, pt. 2, p. 321 (no local-
ity given): Lamarck 1816, Encyclopédie Méthodique, Tableau Mollusques et Polypes Divers, p. 7, Explana-
tion of pl. 430, fig. 5 [error for fig. 4]; Deshayes 1832, ibid., Vers 2, pt. 2, p. 156 (Sénégal and the Antilles).

Pugilina fasciata Schumacher 1817, Essai d’un Nouveau Systéme des Habitations des Vers Testacés, p. 216.

Pyrula morio Linné, Reeve 1847, Conchologia Iconica 4, Pyrula, plate 1, fig. 3.

Melongena morio Linné, Tryon 1881, Manual of Conchology (1) 3, p. 111, pl. 43, figs. 228-229.

Semifusus morio Linné, Dautzenberg 1910, Actes Société Linnéenne de Bordeaux 64, p. 204 (p. 48 of the
reprint).

Melongena (Pugilina) morio Linné, Dautzenberg 1921, Revue Zoologique Africaine 9, p. 112.

Description. Shell reaching 166 mm. (about 64 inches) in length, solid and usually
bluntly spinose. Color a dark chocolate brown, with usually one or more narrow bands
of light yellowish-brown at the periphery. Whorls 9 to 10 and moderately convex. Spire
extended and cast at an angle of about 55°. Aperture narrow, subquadrate and length-
ened below into asomewhat broadened siphonal canal. Anal notch small but well devel-
oped. Parietal wall heavily glazed, usually chocolate brown in color; occasionally it
becomes somewhat lighter in color on the lower columellar area. Outer lip thin but strong
and usually finely crenulated. Columella heavy, fairly broad and twisted. Umbilicus
completely closed. Suture slightly impressed. Sculpture consisting of several blunt spines
which margin the outer edge of the shoulder. Young specimens and the early whorls of
old specimens are axially costate. Occasionally the blunt spines are absent or greatly re-
duced on adult specimens. Spiral sculpture consisting of numerous and fine ridges which
are quite apparent on the early whorls but later become somewhat flattened and incon-
spicuous on the body whorl. In addition, there are several well-marked spiral ridges on
the inner wall of the outer lip. Operculum unguiculate with an apical nucleus and with
fine concentric ridges. Periostracum dull, dark brown in color, thick, somewhat deciduous
and marked with fine axial growth lines. It is thick enough to hide all color of the shell.

length width
166 mm. 80mm. Banana, Belgian Congo
ise 94 Carenage, Trinidad, Lesser Antilles
113 59 Trinidad

* Spire broken

Types. According to Hanley 1855, p. 800, the type specimen is in the Linnean Col-
lection. Linné referred to Bonanni 1684, Recreatio, Rome, plate 357. In the twelfth
edition of the Systema Naturae, Linné referred to Adanson 1757, Histoire Naturelle du
Sénégal, pl. 9, fig. 31. Adanson recorded his species as coming from Gorée Island, Séné-
gal which is here selected as the type locality.

186 JOHNSONIA, No. 35 Pugilina

Remarks. This species has been placed in many different genera since it was originally
described by Linné. We give in our synonymy most of the important name combina-
tions that have appeared.

The typical form of this species as originally described by Linné was without spines.
Later, Lamarck introduced the name coronatus for the spinose form. However, there is
a complete intergradation between these two forms, with specimens of both occurring in
the same locality. Both the spinose and the smooth forms are found in the Western as
well as the Eastern Atlantic. This is another of several species that occur on both sides
of the Atlantic in the tropics.

Range. EasteERN AtrLANtICc: According to Nicklés 1950, p. 101, this species ranges
from Cap Blane, French West Africa, south to Angola, Africa.

Western AtTLANtiIC: Trinidad and south to I]lhado Mel, Est.do Parana, Brasil (Lange
de Morretes 1949, p. 98).

Specimens examined. EASTERN ATLANTIC: SIERRA LEONE: Turtle Island, off Sherbo
Island; Tasso Island, Rokelle Estuary (both T. S. Jones). Liserta: Marshall (MCZ).
Gotp Coast: Accra (MCZ). Cameroons: Longji (MCZ); Batanga (Carnegie Mus.).
Rio Mount: Benito; Corisco Island (both MCZ; Carnegie Mus.). BrELGr1an Conco:
Banana (MCZ).

Western Ariantic: Lesser ANTILLES: Tobago (R. W.Cabot); Balandra, Carenage
and Salena Bay, Trinidad (all H.G. Kugler). Durcu Guiana: Nickerie (D.C. Geijskes).
Brasi.: Salinas, Para (MCZ): Braganga, Para (Carnegie Mus.); Recife, Pernambuco
(MCZ: Carnegie Mus.); Bahia, Bahia (Carnegie Mus.); [ha de Itaparica, Bahia (M.J.
Oliveira): Vit6ria, Espirito Santo (MCZ).

Plate 109. Pugilina morio Linné. Figs. 1-2. Vitéria, Brasil (about 3/4 natural size).

Western Atlantic JOHNSONIA, No. 35 187

The following subgenus, Hemifusus Swainson, is included only to complete the genus.
No species in this subgenus occurs in the Western Atlantic.

Subgenus Hemifusus Stwainson

Hemifusus Swainson 1840, A Treatise on Malacology, London, p. 308 (and 91).
Semifusus Agassiz 1846, Nomenclator Zoologicus, Index, p. 338 [emendation for Hemifusus Swainson ].

Type species, Fusus colosseus Lamarck, subsequent designation, Gray 1847.

Shell attenuate, relatively thin, having generally strong shoulder nodules and posses-
sing a relatively strong sculpture of spiral ridges or threads.

Notes

Pyrula melongena mutica Sowerby 1825, Catalogue of the Shells in the collection of the Earl of Tankerville,
London, p. 62 [nomen nudum].

Fusus pyruloides DeKay 1843, Natural History of New York, Mollusca, Albany, New York, p. 147, pl. 9,
fig. 191 (Ship’s bottom, New York Harbor, believed to have come from a southern port).

We believe this to be a young Thais, perhaps 7’. haemastoma Linné and not a Melon-
gena as stated by Tryon (1881, Manual of Conchology (1), 3, p. 229).

REFERENCES
Abbott, R.T. 1948, Notes on the anatomy of a giant subspecies of Melongena corona from Florida. Shell
Notes, Lantana, Florida 2, pp. 21-24, pls. 1-5.

Baker, F.C. 1891, Notes on a Collection of Shells from Southern Mexico. Proceedings Academy Natural
Sciences, Philadelphia, p. 50.

Bayer, C. 1952, Catalogue of the Genera Melongena and Semifusus. Zoologische Mededelingen, Rijksmu-
seum van Natuurlijke Historie te Leiden 31, no. 25, pp. 265-299.

Dall, W. H. 1889, A Preliminary Catalogue of the Shell-bearing Marine Mollusks and Brachiopods of the
Southeastern Coast of the United States. United States National Museum, Bulletin no. 37, pp. 1-221.
Dall, W. H. 1890, The Tertiary Fauna of Florida. Transactions of the Wagner Free Institute of Science,
Philadelphia 3, pp. 118-122, pls. 8 and 9.

Dollfus, Gustave F. 1887, Une Coquille Remarquable des Faluns de ]’Anjou. Bulletin Société d’Etudes Sci-
entifiques d’ Angers (n.s.) 17, pp. 25-56, pls. 1-4.

Hanley, S. 1855, Ipsa Linnaei Conchylia, London, p. 298.

Heilprin, A. 1887, Explorations on the West Coast of Florida. Transactions Wagner Free Institute of Science,
Philadelphia 1, pp. 70-71, pl. 1, fig. 3.

Krebs, H. 1864, The West Indian Marine Shells. Falster, Denmark, pp. 1-137; ibid. 1947-1948, a repub-
lication by W.J.Clench, C.G.Aguayo and R.D.Turner in Revista de la Sociedad Malacologica *‘Carlos de
la Torre,’’ Habana, Cuba, volumes 5 and 6.

188 JOHNSONIA, No. 35 Melongena

Lange de Morretes 1949, Ensaio de Catalogo dos Moluscos do Brasil. Arquivos do Museu Paranaense 7, p. 98.
Nicklés, M. 1950, Mollusques Testacés Marins de la Cote occidentale d’Afrique. Manuels Ouest-Africains
Z 101.
2, PD.

Olsson, A. A. and A. Harbison 1953, Pliocene Mollusca of Southern Florida. Academy Natural Sciences,
Philadelphia, Monograph 8, pp. 213-214.

Weisbord, N.E. 1926, Notes on the Marine Mollusks from the Yucatan Peninsula, Mexico. Nautilus 39,
pp. 81-87.

Wilcox, Joseph 1896, Notes on Mollusks of Florida. Nautilus 10, pp. 27-29.

ADDITIONS TO THE PHOLADIDAE— PART II
BY
Ruta D. TurNER

The following additions and corrections to the monograph of the Pholadidae seemed
of sufficient importance to publish at this time rather than wait until the Review Num-
ber which will come at the close of Volume 3.

Genus Lignopholas Turner
Johnsonia 3, no. 34, p. 98.

Through an unfortunate omission the citation of a type species for this genus was not
included in the original description. It is Lignopholas clappi Turner.

Jouannetia (Pholadopsis) quillingi T'wrner
Johnsonia 3, no. 34, pp. 139-140.

Since the publication of Johnsonia, no. 84, two additional records for this species have
been received. These extend the range of the species from off Naples, Florida to off Port
Isabel, Texas. Both lots were obtained by shrimp boats. One dead specimen was found
in the mud, while the others were taken from soft rock brought up by the dredge.

Specimens examined: Texas: 40 miles north of Port Isabel in 11-14 fathoms (‘T. Pul-
ley): East-southeast of Port Isabel in 19-20 fathoms (H. Lee).

JOHNSONIA

Published by
THE DEPARTMENT OF MOLLUSKS

Museum of Comparative Zoélogy, Harvard University

Cambridge, Massachusetts

DECEMBER 20, 1957 CYMATUDAE VOL. 3, NO. 36

THE FAMILY CYMATIIDAE IN THE

WESTERN ATLANTIC
BY
WILLIAM J. CLENCH AND Ruta D. TurNER

Most species in the family Cymatiidae are referred to as triton shells. They are wide-
ranging in all tropical seas, a limited few reaching temperate waters. The family contains
more than 100 species, though only a few of these occur in the Western Atlantic. Most
of our Western Atlantic species are identical with species occurring in the Eastern At-
lantic and Indo-Pacific provinces. No other family so far studied in this series of mono-
graphs has had so many species which are so widely distributed. Probably all have a
fairly long veliger or free-swimming stage. This has, of course, aided these animals in
their dispersal; but it is not the whole answer as many other mollusks with equally long
pelagic life have failed to become as widely distributed.

The fossil record indicates that the family was established in the early Tertiary, so that
there has been a time factor of considerable importance. The family probably had its
origin in the Indo-Pacific, and during the existence of the Tethys Sea several species were
able to migrate into the Atlantic Ocean. It would also appear that many specific ele-
ments in this family became stabilized fairly early and have since maintained themselves

ALAS AAS

1 mm,

SANA
\AN\\

Plate 110. Jaws. Fig. 1. Cymatium femorale Linné. Oranjestad, Aruba, Dutch West Indies. Fig. 2. Cyma-
tium gemmatum Reeve. Off Palm Beach, Florida. Fig. 3. Cymatium rubeculum occidentale Clench and Turner.
Carboneras, Rio Camarioca, Matanzas, Cuba. Fig. 4. Cymatium parthenopeum von Salis. Rio de Janeiro,
Brasil.

189

190 JOHNSONIA, No. 36 Cymatiidae

with but little change in the morphology of the shell. This is not without precedent in
other families, such as the Tonnidae, with Jonna galea Linné occurring as it does in the
Eastern and Western Atlantic and in the Indo-Pacific. Examples of this sort are, how-
ever, rare and the present family is quite an exception.

The embryonic shell of all species contained in this family is very different from the
adult shell. The embryonic shell is smooth or only feebly sculptured; it is amber or
brownish in color and the whorls are only slightly convex. The change from this embry-
onic state into the young adult is exceedingly abrupt. Strongly marked sculpture ap-
pears in the form of ridges which are both spiral and axial, and knobs develop where
these ridges cross. The larval or veliger stage may be long, lasting perhaps up to four
weeks, so that dispersal by oceanic currents may play a very important part in their dis-
tribution. The veliger larvae of Cymatium pileare Linné and C. chlorostomum Lamarck
(=C. nicobaricum Réding) have been figured by M. Lebour (1945, pp. 476-477). The
four velar lobes are amazingly long and narrow, being about four times the length of the
shell.

The feeding habits of most species are completely unknown; probably all are preda-
tory, feeding mainly on other mollusks. The senior author found Cymatium femorale
Linné feeding on Laevicardium laevigatum Linné at Savannah Sound, Eleuthera Island,
Bahama Islands.

Plate 111. Opercula. Figs. 1-2. Charonia variegata Lamarck. Molasses Reef, Key Largo, Florida. Figs. 3-4,
Cymatium caribbaeum Clench and Turner. Bear Cut, Miami, Florida. Figs. 5-6. Cymatium nicobaricum Réd-
ing. Molasses Reef, Key Largo, Florida. Figs. 7-8. Cymatium poulsenii Mérch. (Young specimen). Off
Tampico, Mexico. Figs. 9-10. Cymatium labiosum Wood. From an ocean buoy off Cape Romain, South
Carolina.

Western Atlantic JOHNSONIA, No. 36

The jaws consist of two subtriangular and very thin chitinous plates which have n ween
merous longitudinal rows of scales. On the basis of the differences shown in the few jaw’
that we have been able to examine, these may prove to have as much taxonomic value
as the opercula. he jaws are lateral and appear to be structures which aid in opening
the proboscis during feeding. They do not seem to be strong enough to be used as rasp-
ing or grasping organs. The radula, however, is fairly large and strong.

There is a surprising uniformity in the radulae of the various species in the genus
Cymatium. It would appear that, regardless of what modifications have taken place in
the morphology of the shell, the embryonic whorls and the opercula, the radula has re-
mained relatively unchanged. The radula in the genus Charonia, however, is quite dis-
tinct from that of Cymatium (see Plate 113).

The opercula of the several species considered in Cymatium show some important dif-
ferences. In the species caribbaeum, nicobaricum, poulseni and labiosum, the nucleus is
eccentric (Plate 111) while in the remaining species in this genus the nucleus is terminal.
The nucleus of the operculum of Charonia variegata is almost central.

The position of the nucleus of the operculum appears to us to be an important charac-
ter in the classification of the several subgenera. This character emphasizes the relation-
ships between species which are sometimes difficult to see when dealing only with the
several characters of the shell.

From a review of the literature concerning this family, there appears to be no real
agreement among the various authors concerning the limits of the genus Cymatiuwm and

Plate 112. Opercula. Figs. 1-2. Cymatium pileare Linné. Varadero Beach, Matanzas Province, Cuba. Figs.
3-4. Cymatium krebsii Mérch. Sombrero Key Light, Lower Keys, Florida, in 40 fathoms. Figs. 5-6. Cyma-
tium muricinum Réding. Fish Point, Guantanamo, Cuba. Figs. 7-8. Cymatium parthenopeum von Salis. Rio
de Janeiro, Brasil. Figs. 9-10. Cymatium femorale Linné. Bear Cut, Miami, Florida.

192 JOHNSONIA, No. 36 Cymatiidae

its various subgenera. We may be too conservative in considering the various species
groups as only subgenera. However, we still lack so much important data regarding
these animals, such as their life history, ecology and soft anatomy, that a conservative
stand is perhaps best at this time. It must be understood, however, that the various sub-
genera are not at all of equal value. Relationships between the groups are at different
levels. They should be considered convenient categories in our system of classification,
subject to change as more data become available.

Plate 113. Radulae. Fig. 1. Charonia variegata Lamarck. Jérémie, Haiti. Fig. 2. Cymatium poulsenit Mérch.
Tampico, Mexico. Fig. 3. Cymatium caribbaeum Clench and Turner. Bear Cut, Miami, Florida. Fig. 4.
Cymatium nicobaricum Riding. Long Reef, off Elliott Key, Florida. Fig. 5. Cymatium rubeculum occidentale
Clench and Turner. Carboneras, Rio Camarioca, Matanzas, Cuba. Fig. 6. Cymatium gemmatum Reeve. Off
Palm Beach, Florida. Fig. 7. Cymatium pileare Linné. Mauritius. Fig. 8. Cymatium muricinum Réding.
Amboyna Island, Molucca Islands. Fig. 9. Cymatium parthenopeum von Salis. Koka Shima, Japan. Fig. 10.
Cymatium parthenopeum von Salis. Rio de Janeiro, Brasil. Fig. 11. Cymatium femorale Linné. Oranjestad,
Aruba, Dutch West Indies.

All drawings were made with the aid of a camera lucida at a magnification of 100x and the plate is re-
duced to about 50x.

ACKNOWLEDGEMENTS

The loan of material in Cymatium has perhaps been more important to us than in any
other group we have considered in Johnsonia. Most species in this group are quite rare
and it was only by borrowing material widely that we could obtain an understanding of
many of the species. Most of the borrowed material has been recorded under ‘‘Speci- :
mens examined’” and many of the specimens were used in illustrating this paper. A |

Western Atlantic JOHNSONIA, No. 36 193

number of the people listed below also contributed specimens to our collection. We are
most grateful to all for their help. Thanks are extended to R. T. Abbott, Academy of
Natural Sciences, Philadelphia; C.G. Aguayo, Museo Poey, Universidad de la Habana:
E. P. Chace, San Diego Museum of Natural History; R. M. DeWitt, Florida State
Museum (referred to in the text as FSM); L. G. Hertlein, California Academy of
Sciences; M. Keen, Stanford University: H. de Souza Lopes, Inst. Oswaldo Cruz, Rio
de Janeiro, Brasil; H. A. Rehder, United States National Museum; G. L. Voss, Marine
Laboratory, University of Miami; G. Warmke, Institute of Marine Biology, Univer-
sity of Puerto Rico. The following private collectors have also been most helpful in
loaning material and we wish to thank K. Anderson; M. R. Branham; C. J. Finlay;
E. Grigg; K. Johnstone; T. McGinty; R. Merrill; W. Old, Jr.; A. Phares; R.
Robertson: J. S. Schwengel; G. Usticke and R. Walker.

Genus Charonia Giste/

Tritonium Réding 1798, Museum Boltenianum, p. 125 (type species, 7. tritonis Linné | = Murex tritonis
Linné], by tautonymy); non Tritonium O. F. Miller 1776.

Triton Denys de Montfort 1810, Conchyliologie Systématique 2, p. 587 (type species, Triton tritonis Linné
[= Murex tritonis Linné], original designation), non Triton Linné 1758 [Crustacea]; non Triton Laurenti
1768 [Reptilia]; non Triton Fleming 1828 [= Tritonalia Fleming 1828}.

Tritonium “Montfort”? Bowdich 1822, Elements of Conchology, pt. 1, p. 36 [emendation of Triton Denys de
Montfort 1810].

Tritonia Bowdich 1822, Elements of Conchology, pt. 1, caption of plate 10, fig. 4 [error for Tritonium
‘Montfort’ Bowdich 1822]; non Tritonia Cuvier 1798.

Charonia Gistel 1848, Naturgeschichte des Thierreichs fiir héhere Schulen, p. 170, no. 24 [new name for
Tritonium ‘Cuvier’ Réding 1798, non Miller 1776].

Nyctilochus Gistel 1848, Naturgeschichte des Thierreichs fiir héhere Schulen, p. xi [mew name for T'riton
“Broderip’ Montfort 1810; non Laurenti 1768 (Reptilia) ].

Charonis ‘Gistel? Mérch 1877, Malakozoologische Blatter 24, p. 26 [error for Charonia Gistel 1848].

Buccinatorium Mérch 1877, Malakozoologische Blitter 24, p. 26 (type species, Triton nobile Conrad | = T.
variegatum Lamarck | here designated).

Tritonellium ‘Valenciennes’ Mérch 1877, Malakozoologische Blatter 24, p. 25; non Tritonellium Valen-
ciennes 1858.

Septa ‘Perry’ Dall 1904, Smithsonian Misc. Coll’n. 47, p. 134; non Septa Perry 1810 in Arcana.

Eutritonium Cossmann 1904, Essais Paléoconchologie Comparée 6, p. 123. [According to J. Thiele 1929.
We have not seen this publication. |

Eutriton Dautzenberg 1907, Journal de Conchyliologie 55, p. 146 [error for Eutritonium Cossmann 1904].

Type species, T'ritonium tritonis Linné | = Murex tritonis Linné] by tautonymy.

Shells medium to very large in size, some specimens reaching about 430 mm. (17
inches) in length, attenuate, imperforate or only minutely rimate and solid in structure.
Color of dark browns or reddish-browns, usually in marbled patterns. Sculpture generally
consisting of spiral ridges and finer threads, the ridges with or without definite nodules.
Varices present, but apparently produced irregularly. Aperture large and usually flaring,
with sculpture on both inner and outer lips. Siphonal canal distinct and short.

Charonia variegata Lamarck
Plate 111, figs. 1-2; Plate 113, fig. 1; Plate 114, figs. 1-2

Tritonium marmoratum Link 1807, Beschreibung der Naturalien-Sammlung der Universitit zu Rostock, p.
122 [in part, Chemnitz’ reference only; see Remarks below].

194 JOHNSONIA, No. 36 Charonia

Triton variegatum Lamarck 1816, Tableau Encyclopédique et Méthodique, Liste, p. 5, Atlas 3, pl. 421, fig.
2a-b (no locality given); Lamarck 1822, Animaux sans Vertébres 7, p. 178 [in part]; De Blainville 1825,
Manuel de Malacologie, p. 399, pl. 18, figs. 3-3a; Kiener 1842, Iconographie des Coquilles Vivantes Triton,
p. 28, pl. 2.

Trilonia atlantica ‘Montfort’ Bowdich 1822, Elements of Conchology, part 1, plate 10, fig. 4 (no locality
given other than that indicated by the name).

Triton variegatus var. B Reeve 1844, Conchologia Iconica 2, Triton, pl. 1, fig. 3a (West Indies).

Triton nobilis Conrad 1848, Proceedings Academy Natural Sciences Philadelphia 4, p. 121 (West Indies);
Conrad 1849, Journal Academy Natural Sciences Philadelphia (2) 1, p. 212.

Triton commutatus ‘Dunker’ Kobelt 1876 [in] Martini and Chemnitz, Systematisches Conchylien-Cabinet
(2) 3, pt. 2, p. 224 [nomen nudum].

Tritonium seguenzae Aradas and Benoit 1871, Atti dell” Accademia Gioenia di Scienze Naturali di Catania
(3) 5, p. 90 (Sicily); W. Kobelt 1889, Iconographie Europadischen Meeresconchylien 2, p. 19, pl. 35, fig. 1;
pl. $6, fie. 13 pl. S7, fig: A.

Description. Shell large, reaching 331 mm. (about 134 inches) in length, attenuate,
rather heavy in structure, imperforate or with a narrow rimation. Color of the early
whorls salmon-pink changing to a mottled coloration brought about by the alternate
chevron-shaped bars of brown and white which follow the spiral cords. Whorls remain-
ing in the adult 11 to 12, convex, the last proportionately larger and generally producing
a pronounced sloping shoulder. Spire extended and very acutely pointed, forming an
angle of 83° to 48°. Aperture subcircular to ovate. Outer lip expanded and margined
with short plicae that are usually grouped in pairs and are opposite the furrows of the
outside sculpture. Between the paired plicae and at the extreme margin of the lip there
is usually a small dull point. The plicae are white and the space between and extending
to the lip edge is a dark chocolate-brown. ‘The intervals between the paired plicae are a
light brownish-ivory in color. Parietal wall with numerous plicae which extend from the
margin of the parietal shield well within the whorl. Actually these plicae have had a con-
tinuous growth from the very early stages but are absorbed as the animal builds the shell
forward. These plicae are irregular as to width and spacing. They are white and the
area between them is a dark chocolate-brown. Columella fairly wide and slightly arched.
Umbilicus, when present, consisting only of a very narrow rimation beneath the parietal
shield. Siphonal canal short and fairly broad. Suture irregular and not indented. Sculp-
ture consisting of numerous flattened spiral ridges, separated by fairly broad furrows, the
shoulder ridge being the largest. At the bottom of the furrows there may be from one
to three spiral threads. Usually the ridges are narrow and small near the suture and again
at the base of the body whorl. At the suture there isa band that is sculptured with short
axial threads. Nuclear whorls five, smooth and pink in color but generally covered with
a fairly heavy brownish periostracum. First four post-nuclear whorls with a spiral sculp-
ture of nodulose ridges with spiral threads between. ‘These whorls are usually salmon-
pink in color. Beyond the fourth post-nuclear whorl the normal adult sculpture begins.
The first varix appears at the end of the first post-nuclear whorl and is repeated about
every three-fourths of a whorl thereafter. Operculum corneous, elliptical in outline, outer
surface with a central nucleus and concentric growth lines. Inner surface with a large
muscle scar, sculptured with numerous, fine, raised and irregularly concentric threads
which at irregular intervals form little double-looped rings. Area beyond the muscle scar
smooth and highly polished.

195

JOHNSONIA, No. 36

Western Atlantic

™ i is mame
omnis
Bee
i=
—

1

on

TW

2, Char-

Fig. 3. Charonia tri-

Plate 114. Fig. 1. Charonia variegata Lamarck. Bimini Islands, Bahama Islands (about 3/5x). Fig.

onia variegata Lamarck. Cuesco Beach, Guantanamo, Cuba (embryonic shell, 5.6x).

tonis Linné. Ryukyu Islands, Japan (about 2/3x).

196 JOHNSONIA, No. 36 Charonia

length width

331 mm. 168 mm. St. Thomas, Virgin Islands

280 145.5 Lectotype of C. nobilis Conrad

255 113 Santa Barbara de Samana, Santo Domingo, Hispaniola

238 121 Alice Town, North Bimini, Bimini Islands, Bahama Islands

235 120 Sandy Cay, 4 mi. NW of Hope Town, Great Abaco, Bahama Islands

Types. The lectotype of 7. nobilis Conrad is in the Academy of Natural Sciences
Philadelphia, no. 42537 from the West Indies. According to Kiener the type of 7.
variegata Lamarck is in the Paris Museum. We here restrict the type locality to St.
Thomas, Virgin Islands.

Remarks. This is one of the largest of our Western Atlantic gastropods. Though not
rare, it does not seem to be common at any one locality.

It is very close in relationship to Charonia tritonis Linné (Plate 114, fig. 3), a wide-
ranging species of the Indo-Pacific region but it differs from that species by never attain-
ing the large size of tritonis. In addition, C. variegata differs by having a fairly well
developed shoulder but a narrower and less flaring lip in the adult. The plicae on the
outer lip of tritonis are proportionately much flatter and in many cases they are absent
or are indicated only by the color markings. ‘The plicae on the outer lip of varvegata are
always very well developed and are generally grouped in pairs. The plicae on the parie-
tal wall of tritonis are much broader than those of variegata, so they limit definitely the
amount of brown coloration on the columella.

There has been a great deal of confusion regarding the names to be employed for the
Atlantic and Indo-Pacific species in this complex. Most of the early descriptions were
composites, and many references by Linné, Gmelin and others were given to either one
or the other indiscriminately. In Linné’s original description of tritonzs his citations were
to Bonnani, Rumphius and Gualtieri. The first two refer to the Indo-Pacific species,
tritonis, and the last is to the Atlantic species. We here select the figure in Bonnani
1684, Recreatio, fig. 188, to be the type figure of tritonis Linné. In 1807, Link (Joc. cit.)
introduced Tritonium marmoratum as a substitute name for Murea tritonis Linné. Link’s
first reference was to Gmelin 1791, Systema Naturae, ed. 13 and his second was to Chem-
nitz 1780, Conchylien-Cabinet. We here restrict Link’s name to Gmelin’s reference to
Bonnani, Recreatio, fig. 188, so that both names (fritonts Linné and marmoratum Link)
are now based upon the same figure.

So far as we have been able to determine, the earliest valid name for the Western
Atlantic species is T'riton variegatum Lamarck. This species has usually been dated from
Lamarck’s description in the Animaux sans Vertébres 1822. However, Lamarck origi-
nally named this species in the Tableau Encyclopédique et Méthodique in 1816, referring
to the figure in the Atlas. This figure is very definitely of the Atlantic species. Later,
in 1822, as stated above, Lamarck described this species and gave several references which
included both the Indo-Pacific and the Atlantic forms. Because of this later description,
Lamarck’s variegata has been considered a synonym of C. tritonis Linné, as his earlier
use of the name in 1816 had been completely overlooked.

This species is generally found about rocky reefs below low water. Specimens taken
from fish traps that had been placed in 60 to 120 fathoms are very much lighter in struc-
ture and somewhat smoother. Mr. C. J. Finlay reports that these specimens were not
alive but were inhabited by hermit crabs.

Western Atlantic JOHNSONIA, No. 36 197

Range. Easrern AtiLantic: Mediterranean Sea, Cape Verde Islands, the Canary
Islands and St. Helena. WesrerN AtrLaNtric: Bermuda (A. J. Peile), the Bahama Is-
lands, the Lower Florida Keys, the West Indies and from central Mexico south to
Santos, Estado Sio Paulo, Brasil.

Specimens examined. WersSTERN ATLANTIC. FLORIDA: Molasses Reef, Key Largo
(MCZ; R.Work); Key West (ANSP); Tortugas (MCZ). Bermupa: Castle Harbour
(H. B. Moore—dredged and dead). BaHama IsLanps: Elbow Cay; Sandy Cay, 4 miles
NW of Hope Town, both Great Abaco (both R. Robertson); Alice Town, North Bimini
Island, Bimini Islands (MCZ); Pigeon Cays, Andros (J. Schwengel); Bullocks Har-
bour, Great Harbour Cay, Berry Islands; Dicks Point, Nassau, New Providence; The
Current, Eleuthera Island (all G. Kline); Spanish Wells, northern Eleuthera (J. Schwen-
gel); Governors Harbour, Eleuthera; Little San Salvador Island; Arthurs Town, Cat
Island (all MCZ); Channel Cay, Great Exuma Island (G. Kline); Clarence Town, Long
Island; Abraham’s Bay, Mariguana Island; Matthew Town, Great Inagua (all MCZ).
Cusa: Cabanas Bay, Pinar del Rio(USNM); Matanzas Bay, Matanzas in 90-120 fath-
oms (C. J. Finlay); Cayo Francés, Caibarién, Las Villas (P. Bermudez); Cayo Largo,
Banco de los Jardines, Las Villas (ANSP); Cayo Maja Figuro, off Punta Alegre, Cama-
giiey (R. Humes); off Gibara, Oriente in 60-80 fathoms (C. J. Finlay); Cuesco Beach,
Guantanamo Naval Base, Oriente (MCZ). Hispaniota: Monte Cristi; Blanco, 30 km.
NW of Puerto Plata; Puerto Plata; Puerto Sosta; Santa Barbara de Samana, all Santo
Domingo (all MCZ); Saltrou, Dept. de Quest; Roche a Bateau, Dept. du Sud; Ile
de la Gonave, all Haiti (all USNM); Jérémie, Dept. du Sud, Haiti (MCZ). Jamaica:
St. Anns Bay: near Buff Bay, Portland; Bull Bay, St. Andrews (all USNM). PuErro
Rico: Arecibo (G. Warmke); Mona Island (A. Phares); Rineén Light, Rincon (G.
Warmke; MCZ); reef 6 miles off Mayagiiez (K. Yates); Mayagiiez (G. Warmke);
Punta Cuchara, about 5 mi. SW of Ponce (G. Warmke). VirGin IsLanps: St. Thomas
(ANSP; USNM): The Baths, Virgin Gorda(M. Dewey). Lesser ANTILLES: St. Kitts
(MCZ); Barbados (MCZ; USNM): St. Vincent (USNM); Grenada (IX. Anderson) ;
Bucco Reef, Tobago (MCZ; ANSP); Cyril Bay, Trinidad (H.G. Kugler). CaripBEan
IsLANDs: Swan Island (MCZ); Grand Cayman, Cayman Islands (ANSP). Mexico:
Lobos Island, near Tampico; Tampico, both Tamaulipas (both 'T. Pulley): Veracruz,
Veracruz (M. E. Bourgeois; ANSP); Isla Mujeres, Yucatan (C.G. Aguayo). Panama:
Col6bn (MCZ; USNM). Cotomsia: near Cartagena (USNM). VENEZUELA: Tucacos
Bay, Estado Faleén (H.G. Kugler). Brasit: Santos, Estado Sao Paulo (ANSP).

EASTERN ATLANTIC. MEDITERRANEAN IsLanps: Off Old Limassol, Cyprus (J. K.
Howard). LreBanon: Beirut (USNM).

Genus Cymatium Réding

Cymatium Réding 1798, Museum Boltenianum, p. 129.

Lotorium Denys de Montfort 1810, Conchyliologie Systématique 2, p. 583 (type species, Lotorium lotor
Montfort [= Murex femorale Linné] original designation); non Lotorium Pusch 1837).

Currus Lesson 1842, L’Echo du Monde Savant (2) 6, Col. 65 [reference from Dall 1904, p. 139. We have
not seen this publication ].

Luterium Herrmannsen 1846, Indicis Generum Malacozoorum 1, p. 632. [Emendation of Lotorium Denys

de Montfort. |

Type species, Murex femorale Linné, subsequent designation, Dall 1904.

198 JOHNSONIA, No. 36 Cymatium

Shells ranging in size from about 25 mm. (1 inch) to 212 mm. (about 8 inches) in
length. Generally brown in color with bands of darker brown or red-brown. Many spe-
cies show remnants of banding on the varices. Nearly all known species have one to six
varices. They are generally sculptured with spiral cords which may or may not be beaded
and, in many cases, the cords are knobbed. Siphonal canal generally fairly short and
turned upward. Inside of outer lip usually with fine to coarse denticles. Parietal area
generally having numerous lamellae. Periostracum generally heavy and produced in axial
fringed blades. Operculum subcircular, sculptured with concentric growth lines and with
an eccentric or terminal nucleus. Embryonic whorls fairly large, generally persistent and
usually smooth or sculptured with microscopic axial striae.

All members of this genus occur only in the tropics or in the warmer portions of the
temperate zone. They generally are found from the low water line out to depths of a
little over 100 fathoms.

The genus Cymatium is composed of a number of subgenera many of which have been
raised by various authors to generic rank. However, considering this genus from a world
point of view, many of the characters used for separating the groups overlap. Conse-
quently, it is impossible to assign clear cut definitions and it is often difficult to place
species in the proper subgenus.

Subgenus Linatella Gray

Linatella Gray 1857, Guide to the Systematic Distribution of Mollusca in the British Museum, p. 39.
Zinatella Cossmann 1903, Essais de Paléoconchologie Comparée 5, p. 86 [error for Linatella Gray ].

Type species, L. cingulata Lamarck [= C. cynocephalum Lamarck] monotypic.

Shell medium to small in size and generally uniformly light yellowish brown in color
and, in some, spirally banded with dark brown, particularly on the spiral cords. Sculp-
ture consisting of numerous low and occasionally knobbed spiral cords. Only a few
specimens produce more than one lip varix. Siphonal canal short and turned upward.
Aperture subcircular. Outer lip with fine to fairly coarse denticles. Parietal area with or
without lamellae. Periostracum rather thin, deciduous and produced in numerous low
axial blades which are minutely fringed. Operculum subcircular, with an eccentric nu-
cleus and sculptured with concentric growth lines.

Cymatium (Linatella) poulsenii Mérch
Plate 111, figs. 7-8; Plate 113, fig:2; Plate 114, tgs. 1-3

Triton (Linatella) poulsenii Mérch 1877, Malakozoologische Blatter 24, p. 33 (Curacao and Porto Cabello
[Venezuela] ).

Fusus cutaceus Lamarck 1816, Tableau Encyclopédique et Méthodique 3, pl. 427, fig. 4a—b; Liste, p. 6;
non T'rifon culaceus Lamarck‘ 1816, ibid. pl. 414, fig. 2a—-b, Liste, p. 4 which is Cymatium cutaceum Linné.

Cymatium cingulatum peninsulum M. Smith 1937, East Coast Marine Shells, p. 113 (Lake Worth, Florida).

Description. Shell moderate in size, reaching 75 mm. (about 3 inches) in length, rather
thin in structure, imperforate or nearly so, and spirally sculptured. Color light straw.

leny- * . - . . r .
rhis species has often been credited to Lamarck; but in 1822, Animaux sans Vertébres 7, p. 188, La-
marek definitely gives credit to Linné and refers to the same figures in Seba. This is a Mediterranean species.

Western Atlantic JOHNSONIA, No. 36 199

yellow toa medium brown and occasionally banded with darker brown. Specimens occur
rarely with axial stripes of darker brown. Post-embryonic whorls four, convex, with the
body whorl slightly shouldered. Spire moderately extended and produced at an angle of
about 75°. Aperture subelliptical in outline, the outer lip crenulated and slightly ex-
panded. Inner lip with a light glaze on the body whorl and a much heavier glaze over
the columella. Siphonal canal variable, moderately long and curved upward. Columella
arched inward, its base continuing as the parietal margin of the siphonal canal. Suture
slightly indented. Sculpture consisting of 18 to 20 major flattened spiral cords which are
somewhat variable in height and width. In some specimens finer spiral cords may be
present between the major cords. The shoulder cord of occasional specimens may be
slightly beaded. Axial sculpture consisting of fine growth lines, with occasional speci-
mens having a thin blade-like varix. Periostracum thin, usually deciduous and consisting
of numerous fine axial blades from which extend short hair-like processes. Operculum
thin, corneous, subcircular in outline, with concentric growth lines and an eccentric nu-
cleus. Embryonic whorls 38 to 34, smooth, yellowish horn in color, narrow, extended,
and set off from the later whorls.

length width
75 mm. 43 mm. Lake Worth, Florida
65 4.5.5 off Port Isabel, Texas
58 33.5 off Laguna Madre, Tamaulipas, Mexico

Types. The holotype of 7°. poulseni Mérch is probably in the Universitetets Zoolo-
giske Museum, Copenhagen, Denmark. The location of the holotype of F. eutaceus
Lamarck is unknown tous. The holotype of Cymatium cingulatum peninsulum M.Smith
is in the Florida State Museum, Gainesville, Florida.

“ +
ok F
ey —_

ot?

wa ere
pamela
See a ‘adlineel “er

Algcepree

Plate 115. Cymatium poulsenii Mérch. Fig. 1. South end of Lake Worth, Palm Beach Co., Florida (slightly
enlarged). Fig. 2. Off Laguna Madre, Tamaulipas, Mexico (1.24x), Fig. 3. Off Laguna Madre, Tamau-
lipas, Mexico (17x).

200 JOHNSONIA, No. 36 Cymatium

Remarks. This species is among the rarest of our Western Atlantic cymatiids. It has
a fairly wide range throughout the Gulf of Mexico and the Caribbean Sea and, so far as
now known, it lives well below low water line. Our few depth records range from 12
to 209 fathoms. The specimens from off the Barbados in 209 fathoms were apparently
dead when collected and they may be advectitious at this depth.

This species is closely related to Cymatium cynocephalum Lamarck’ from the Eastern
Pacific. It differs by being much smaller and in having the spire far less extended.

Range. From Lake Worth, Florida south through the West Indies, and from west
Florida to Texas, Mexico and south to Venezuela.

Specimens examined. FioripA: Palm Beach in 14 fathoms (J. Schwengel); Lake
Worth (FSM: T. McGinty): Boynton Beach; off Delray Beach (both T. McGinty);
Bear Cut, Crandon Park, Biscayne Bay (R. Merrill); Bahia Honda Key; off Key West
in 11 fathoms (both H. and K. Johnstone): off Tortugas, from shrimp boat in 18-20
fathoms (R. Merrill: T. McGinty); Naples (ANSP). Texas: 30 miles N of Port Isabel
(H. Hildebrand); off Port Isabel (W. C. Frisby; H. and Kk. Johnstone: 'T. McGinty).
Mexico: off Laguna Madre, Tamaulipas in 12 to 14 fathoms (C. L. Branch); Tampico,
Tamaulipas (T. Pulley); about 70 miles NNE of Tampico, Tamaulipas (W. C. Frisby);
15 miles N of Tecolutla, Veracruz (T. Pulley). Cusa: Puerto Esperanza, Pinar del Rio
(C. G. Aguayo). Virein Istanps: St. Thomas (ANSP). Lesser ANTILLES: off Bar-
bados, Blake station 274 (N. Lat. 13°00/50’; W. Long. 59°36’) in 209 fathoms (MCZ).
VENEZUELA: Cubagua Island off Cumana (MCZ).

Subgenus Cabestana Roding

Cabestana Réding 1798, Museum Boltenianum, p. 130.

Aquillus Denys de Montfort 1810, Conchyliologie Systématique 2, p. 579 (type species, Aquillus cutaceus
[ = Murex cutaceus Linné] original designation).

Dolarium Schlueter 1838, Systematisches Verzeichniss meiner Conchyliensammlung, Halle, p. 20 (type
species, Murex caduceus {error for cutaceus Linné], monotypic).

Aquilus Mirch 1852, Catalogus Conchyliorum Comes de Yoldi, p. 108 (error for Agquillus Montfort).

Neptunella Gray 1853 [1854] Proceedings Zoological Society London, p. 38 (type species, Murer cutaceus
Linné, monotypic); non Neptunella Meek 1864; non Verrill 1873.

Tritonicus Dall 1904, Smithsonian Miscellaneous Collections 47, p. 134 (type species, Triton loroisi Petit
de la Saussaye | =C. labiosum Wood] original designation).

Turritriton Dall 1904, Smithsonian Miscellaneous Collections 47, p. 133 ie pe species, Triton gibbosus Brod-
erip, original designation).

Particymatium Iredale 1936, Records Australian Museum 19, p. 307 (type species, Triton strangei Adams
and Angas, monotypic).

Cabestanimorpha Iredale 1936, Records Australian Museum 19, p. 307 (type species, Triton exaratum Reeve,
monotypic).

Cymatilesta Iredale 1936, Records Australian Museum 19, p. 307 (type species, Triton spengleri Lamarck,
monotypic).

Parlicymatium ‘lredale’ Wenz 1941, Handbuch der Palaozoologie 6, pt. 1, p. 1063 [error for Particymatium
Iredale].

Type species, Murex cutaceus Linné, subsequent designation, Dall 1904.

ae : s . - :
his is an earlier name for Cymatium cingulatum Lamarck. See remarks under C. caribbaeum Clench and

furner, new name, regarding the use of the name cynocephalum Lamarck.

———<—

Western Atlantic JOHNSONIA, No. 36 201

Shells relatively small to medium in size and generally brownish in color. Sculpture
consisting of a few varices and rather strongly developed, knobbed, axial costae. Many
species in this subgenus have very well developed spiral cords. Whorls with a broad
shoulder. Siphonal canal short. Shells usually rimately umbilicate. Periostracum thin.
Operculum unguiculate, with a marginal nucleus.

There are two species in this subgenus in the Western Atlantic.

Cymatium (Cabestana) labiosum Wood
Plate 111, figs. 9-10; Plate 116, fig. 1

Murex labiosus Wood 1828, Index Testaceologicus, Supplement, p. 15, pl. 5, fig. 18 (locality unknown):
non Murex labiosa J. E. Gray 1828; non G. D. Nardo 1847.

Tritonium rutilum Menke 18438, Molluscorum Novae Hollandiae, p. 25 (litore occidentali [shore of western
Australia] ).

Triton loroisi Petit de la Saussaye 1852, Journal de Conchyliologie 3, p. 53, pl. 2, fig. 8 (Guadeloupe
[Lesser Antilles] ).

Triton strangei Adams and Angas, July 1864, Proceedings Zoological Society London, p. 35 (Moreton Bay
[Queensland] Australia); Smith 1878 [1879] Proceedings Zoological Society London, p. 816, pl. 50, fig. 16.

Triton (Gutturnium) orientalis G. and H. Nevill 1874, Journal Asiatic Society Bengal 43, p. 29. Refers to
figure in Reeve, Conchologia Iconica 2, Triton, pl. 11, fig. 38 (Andamans, dredged).

Description. Shell rather small, reaching 29.5 mm. (about 14 inches) in length, solid,
minutely perforate, shouldered and strongly sculptured. Color generally mahogany-
brown to yellowish and occasionally with a white band at the middle of the whorl. In-
terior of aperture white. Post-embryonic whorls 5 and strongly convex. Spire moderately
extended and produced at an angle of about 60°. Aperture subcircular, with the outer
lip greatly thickened when a varix is produced. Parietal lip narrow and thickly glazed.
Outer lip with 6 flattened and rather obscure denticles. Inner lip with very weak lamel-
lae — usually one near the anal canal and two or three near the entrance of the siphonal

Plate 116. Fig. 1. Cymatium labiosum Wood. Carboneras, Rio Camarioca, Matanzas, Cuba .8x). Fig. 2.

(2
Cymatium felipponei von Ihering. Praia de Guarariba, Vitoria, Estado Espirito Santo, Brasil (1.85x).

202 JOHNSONIA, No. 36 Cymatium

canal. Columella slightly arched inwardly with its base continuing into the parietal mar-
gin of the short siphonal canal. Suture slightly indented. Sculpture consisting of 4 to 6
strong, nodulose, spiral cords. In addition, these cords are sculptured with 3 or 4 beaded
threads. Beaded threads are also found in the interspaces between the major cords. From
each knob on the shoulder cord there is a ridge extending to the suture. This ridge con-
tinues axially to the base of the shell. Periostracum straw-yellow in color and consisting
of very low thin axial blades. Operculum unguiculate, with a marginal nucleus and sculp-
tured with concentric growth lines. Embryonic whorls 34, moderately convex, amber in
color and with very fine axial striae.

length width

29.5 mm. 18.5 mm. North Inlet, Lake Worth, Florida

27 18 Carboneras, near Rio Camarioca, Matanzas, Cuba
26.5 16.5 Zanzibar Island

24 15 St. Thomas, Virgin Islands

99.5 ts Funafuti, Ellice Islands

Types. The location of the type of C. labiosum Wood is unknown. It was originally
in the cabinet of a Mrs. John Mawe and according to Sherborn it was purchased by
‘Tennant of the Strand.*’ As the locality was unknown we here designate Guadeloupe,
Lesser Antilles as the type locality. The type of C. strangei Adams and Angas was in the
Angas Collection and is now probably in the British Museum. An idiotype from Angas
is in the Museum of Comparative Zoology. The type locality is Moreton Bay, Queens-
land, Australia. The type of C. loroisi Petit de la Saussaye is in the collection of the
Journal de Conchyliologie: the type locality is Guadeloupe, Lesser Antilles.

Remarks. This is a very distinctive species, not closely related to any other in the
Western Atlantic. It is nearest in relationship to C. felipponet von Ihering from which
it differs in being much smaller and having a more pronounced axial sculpture and a pro-
nounced angle to the whorl shoulder. In the Indo-Pacific there appear to be two or three
fairly closely related species. Cymatium waterhousu Adams and Angas from South Aus-
tralia has a proportionately much larger aperture and C. klenei Sowerby from South
Africa is much more strongly shouldered and sculptured, as well as being a larger and
more elongate species.

This appears to be a rather rare species and many of the specimens in collections are
dead beach shells. Nothing is known concerning depth range or habitat of this species.

Range. WrsterN ArLANtic: From off Cape Romain, South Carolina: Florida; the
Bahamas and south through the Lesser Antilles.

InNpo-Paciric: The Hawaiian Islands, west to the Philippine Islands, south to New
South Wales, Australia and west to East Africa.

Specimens examined. WESTERN ATLANTIC. SourH CAROLINA: From ocean buoy off
Cape Romain (R. Merrill). FLoripa: off Palm Beach in 80 fathoms (T. McGinty); Lake
Worth, Palm Beach County (FSM; J. Schwengel); Boca Raton (T. McGinty); Bear
Cut, Key Biscayne (R. Work); Pelican Shoals, Key West (IT. McGinty); Tortugas
(FSM). Banama Istanps: Eight Mile Rock, Grand Bahama Island (MCZ): Drunken

Western Atlantic JOHNSONIA, No. 36 203

Cays and Cooper Jacks Cays, Great Abaco; 4 miles W of Nassau, New Providence (all
R. Robertson); North Cay, New Providence: off Nassau in 4-6 fathoms; North Bimini
Island, Bimini Islands; Gun Cay, Bimini Islands; Pigeon Cay, Andros (all T. McGinty).
Cupa: Vedado, Habana; Camarioca, Matanzas (both Museo Poey); Carboneras near
Rio Camarioca, Matanzas (C. J. Finlay); Varadero, Matanzas (C. J. Finlay; Museo
Poey); Cayo Santa Maria, off Punta Alegre, Camagiiey (R. Humes); Guantanamo Bay
(MCZ); near Cienfuegos, Las Villas (Museo Poey). Purerro Rico: Aguadilla (A.
Phares); Rincén Light House, Rineén (G. Warmke). VirGin IsLtanps: 1 mile N of
Frederiksted, St. Croix (G. Usticke); St. Thomas, Virgin Islands (MCZ). Lesser
ANTILLES: Tortola (MCZ); St. Kitts (R. W. Jackson).

Inpo-Paciric. ELiice IsLanps: Funafuti(MCZ). Paruiprine IsLanps: Zamboanga,
Mindanao (MCZ). Ausrratia: Port Jackson, New South Wales (MCZ). INponesia:
Manipa Island, Molucea Islands (MCZ).

INpIAN OcEAN IsLANDsS: Ceylon (MCZ); Zanzibar Island (MCZ).

Cymatium (Cabestana) felipponei von Ihering
Plate 116, fig.2

Lotorium felipponei von Ihering 1907, Anales Museo Nacional de Buenos Aires (3) 7, p. 443, pl. 18, fig.
122a—b (Maldonado, Uruguay).

Description. Shell medium in size, reaching 52 mm. (about 2 inches) in length, mod-
erately heavy in structure, imperforate or with a small umbilical chink, and with pro-
nounced sculpture. Color a light reddish brown. Post-embryonic whorls 5, convex and
shouldered. Spire moderately extended and produced at an angle of about 45°. Aper-
ture subelliptical, the outer lip margined with rather strong teeth, the inner lip smooth,
forming a thickened shield over the parietal area. Columella arched inward; siphonal
canal short and curved upward slightly. Axial sculpture consisting of 2 to 4 strong vari-
ces; in addition there are numerous and rather fine incised lines which cut across the
spiral cords. Spiral sculpture consisting of numerous cords which are somewhat irregular
in width, the shoulder cord usually being the largest. Periostracum thin, straw-yellow in
color. Operculum and embryonic whorls unknown.

length width
52 mm. 31mm. Mar del Plata, Argentina (from Carcelles 1944, p. 246)
45 30 Punta del Este, Maldonado, Uruguay
42.5 27) Praia de Guarariba, Vitoria, Estado Espirito Santo, Brasil

Types. According to von Ihering 1907, p. 1, the types of this species are in the Museo
Nacional de Buenos Aires. The type locality is Maldonado, Uruguay.

Remarks. This species appears to be distantly related to Cymatium cutaceum Linné of
the Mediterranean, from which it differs in being less highly sculptured and having only
a slight indication of an umbilicus. See also Remarks under C. labiosum Wood.

This is avery rare species to judge by the few specimens that are known to exist. We
are indebted to Eliseo Duarte of Montevideo, Uruguay for two lots of this species, the
one from Vitoria, Brasil extending the known range of this species about 1400 miles to
the north.

204 JOHNSONIA, No. 36 Cymatium

Range. From Vitoria [Victoria], Brasil south to Puerto Quequen, Argentina.

Specimens examined. Brasiu: Playa de Guarariba, Vitoria, Est. Espirito Santo (EK.
Duarte). Urvucuay: Cabo Santa Maria (A. Carcelles); Punta de] Este, Maldonado
(E. Duarte).

Subgenus Ranularia Schwmacher

Ranularia Schumacher 1817, Essai d’un Nouveau Systéme, p. 253,

Ranula Schumacher 1817, Essai d’un Nouveau Systéme, p. 77

Tritonocauda Dall 1904, Smithsonian Miscellaneous Collections 47, p. 133 (type species, Murex caudatus
Gmelin, original designation). [New name for Ranularia ‘Schumacher’ Fischer, non Ranularia Schumacher. |

Type species, T'ritonium clavator Chemnitz [= Ranularia longirostra Schumacher]
subsequent designation, Herrmannsen, 1847.

Herrmannsen in his Indicis Generum Malacozoorum 2, p. 388, gave the type species
as T'ritonium clavator Chemnitz. However, Schumacher used /ongirostra, with clavator
Chemnitz given as a synonym.

Schumacher made a curious error when he instituted the name Ranula (p. 77) in his
generic system, and later (p. 258), he used Ranularia referring by number back to Ran-
ula. However, his second use carries not only the description but here he cites two spe-
cies. The brief generic diagnosis under Ranula would be unidentifiable standing alone.

The new name T'ritonocauda introduced by Dall for Ranularia Schumacher as used
by Fischer (1884, Manuel de Conchyliologie, p. 655) was unnecessary and partly in error,
for Dall gave as the type, Murea caudatus Gmelin, a name not mentioned by Fischer.

Shell medium in size and generally yellowish to brownish in color. Sculpture consist-
ing of a few to several rounded varices, with most species having strongly developed,
knobbed, axial costae. Spiral sculpture consisting of fairly strong cords which may be
knobbed or beaded. Between these cords there are usually numerous, fine, spiral threads.
Whorls convex. Siphonal canal moderately extended. Aperture subcircular with a mod-
erately thickened parietal wall. Periostracum produced in numerous axial blades which
are fringed with hair-like processes. Operculum subovate, with a submarginal nucleus
located about midway near the parietal border.

Cymatium (Ranularia) caribbaeum, new name
Plate 111, figs. 3-4;° Plate 113, tig. 3; Plate 117) tes, 1-2

Triton cynocephalum ‘Lamarck’ Kiener 1842, Iconographie des Coquilles Vivantes, Triton, p. 3, pl. 12, fig.
1 (Bahia, Brasil); non Triton cynocephalum Lamarck 1816 and 1822.

Cymatium cyanocephalum ‘Lamarck’ Johnson 1934, Proceedings Boston Society Natural History 40, p. 114
[error for cynocephalum Lamarck }.

Description. Shell medium in size, reaching about 84 mm. (about 84 inches) in length,
solid, imperforate and strongly sculptured. Color a light tan to a rich cinnamon brown;
the varices with alternating bands of tan and white. Occasionally specimens are found
in which the white band of the varices extends as a spiral band from varix to varix.
W horls 55 to 6, convex and shouldered. Spire short and broad and produced at an angle
of about 65°. Aperture subcircular; outer lip thickened and having seven rather coarse
denticles. Inner lip consisting of a thickened shield on which there is a series of small

Western Atlantic JOHNSONIA, No. 36 205

plicae set well within the aperture and not extending to the edge of the shield. The area
between the plicae is a dark red-brown. The edge of the lip is usually a light tan or salmon
color. Siphonal canal extended and slightly curved upward and to the left. Columella
thickened and continued into the inner margin of the siphonal canal. Suture slightly in-
dented. Sculpture consisting of about 7 nodulose, spiral cords, with numerous fine spiral
threads between the cords. Axial sculpture consisting of two or three coarsely knobbed
varices, the knobs being formed where the spiral cords cross over the varix. Between
the varices, the spiral cords support from 8 to 18 small knobs. The knobs on the cords
are in axial alignment, but are not noticeably connected. Those on the shoulder cord are
usually the largest. Operculum subovate, sculptured with numerous concentric ridges and
with the nucleus submarginal, about midway on the parietal side. Periostracum usually
deciduous, yellowish to brown in color, thin in texture and consisting of fringed axial
blades. Embryonic shell of 3 to 4 whorls which are amber in color, smooth, moderately
extended and mainly chitinous. The first post-embryonic whorl is calcareous, spirally
ribbed and slightly envelops a part of the embryonic shell.

length width
84 mm. 45mm. off Key West, Florida
67.5 4] Dry Tortugas, Florida
63 32 Puerto Plata, Santo Domingo
60 33.5 Barbados, Lesser Antilles

Types. The holotype of Triton cynocephalum Wiener is probably in the Paris Museum.
The type locality is Bahia, Brasil.

Plate 117. Cymatium caribbaeum Clench and Turner. Fig. 1. Cardenas Keys, Matanzas, Cuba (1.2x). Fig. 2.
Bear Cut, Miami, Florida (1. 16x).

206 JOHNSONIA, No. 36 Cymatium

Remarks. It seems incredible that an error should have persisted so long in the nam-
ing of this species. So far as we can trace it, this error was initiated by L. C. Kiener in
his Iconographie des Coquilles Vivantes. He figured under the name of cynocephalum
Lamarck a shell which is totally different from that originally figured by Lamarck in the
Tableau Encyclopédique et Méthodique. Lamarck’s original figure of cynocephalum(1816)
is of the species generally known today as Cymatium cingulatum of the Panamic Province.
Under the section Notes at the end of this report we give the synonymy of this Eastern
Pacific species and figure a specimen from Bahia Magdalena, Baja California, as well as a
copy of Lamarck’s original figure for comparison.

Cymatium caribbaeum is close in its relationships to C. sarcostomum Reeve, from which
it differs by having many small uniform knobs on the spiral cords. In C. sarcostomum
there are only a few large, irregular knobs. From C. pyrum rehderi Verrill it differs like-
wise in the type of knobs; in addition, the plicae on the parietal wall do not extend to
the outer margin of the narrow parietal shield of caribbaeum, while in rehderi they do
reach to the outer edge.

Cymatium caribbaeum Clench and Turner is confined entirely to the warmer portions
of the Western Atlantic.

Range. Bermuda, southern Florida, the West Indies and Central Mexico south to
Bahia, Brasil.

Specimens examined. FLoripa: Lake Worth (FSM); Bear Cut, Key Biscayne, Miami
(R. Work: G. Voss; R. Merrill); West Summerland Key; Missouri Key (both D. and
N. Schmidt); Pelican Shoals, off Boca Chica Key (MCZ); Key West (MCZ; H. and
K. Johnstone); Tortugas (MCZ; FSM; R. Merrill). BermMupa: Castle Harbour
(dredged dead, H. Moore). BanamMa Istanps: Elbow Cay, Great Abaco (G. and M.
Kline; R. Robertson); Alice Town, Bimini Islands (MCZ); Dicks Point, Nassau, New
Providence (G. and M. Kline); Simms, Long Island; Matthew Town, Great Inagua
(both MCZ). Cusa: Habana, Habana (Museo Poey); Cardenas, Matanzas (J. Finlay);
Cayo Francés, Caibarién, Las Villas (P. J. Bermudez); Cayo Santa Maria, Camagiiey
(R. Humes); Gibara, Oriente (Museo Poey); Caleton de Don Bruno, Cienfuegos, Las
Villas (MCZ); Guantaénamo Bay, Oriente (MCZ; Museo Poey). Jamaica: White
House (J. K. Howard). Hispanioua: Puerto Plata and Santa Barbara de Samana, Santo
Domingo (both MCZ). Purrro Rico: Catano; Rinecén; Punta Algarrobo; Punta
Guanajibo; Playa de Naguabo (all G. Warmke). VirGin Istanps: Virgin Gorda
(M. W. Dewey); St. John (MCZ); St. Croix (R. Walker; FSM; G. Usticke). Lesser
ANTILLES: Barbados (MCZ); Chaguaramas, Trinidad (H. G. Kugler); Mexico: Tam-
pico, Tamaulipas; Veracruz, Veracruz (both T. Pulley). Panama: 2 miles off Colén in
10 fathoms (W. C. Clarke). Brasu.: Itaparica, S40 Salvador, Bahia (H. S. Lopes).

Cymatium (Ranularia) sarcostomum Reeve
Plate 118; fies. 1-5

Triton sarcostoma Reeve, April 1844, Conchologia Iconica 2, Triton, pl. 7, fig. 21a—b (Island of Ticao, Phil-
ippines); Reeve, Dec. 1844, Proceedings Zoological Society London 12, p. 114.

Triton moritinctus Reeve, June 1844, Conchologia Iconica 2, Triton, pl. 13, fig. 49 (Philippine Islands);
Reeve, Dec. 1844, Proceedings Zoological Society London 12, p. 115.

Triton ridleyi Smith 1890, Journal Linnean Society London 20, p. 489, pl. 30, fig. 1 (Fernando Noronha
Island, Brasil).

Western Atlantic JOHNSONIA, No. 36 207

Description. Shell medium in size, reaching 60 mm. (about 22 inches) in length, solid,
imperforate and strongly sculptured. Color light straw-yellow to reddish brown, occa-
sionally banded with white, particularly on the varices. Whorls 5 to 6, convex and
shouldered. Spire short and broad and produced at an angle of about 65°. Aperture sub-
circular. Outer lip thickened and having 7 rather coarse denticles. Inner lip consisting
of a thickened shield on which there is a series of small plicae set well within the aper-
ture and not extending to the edge of the shield. The area between the plicae is usually
a dark red-brown, the edge of the lip white to a light tan. Siphonal canal extended and
slightly curved upward. Columella thickened and continued into the inner margin of
the siphonal canal. Suture slightly indented. Spiral sculpture consisting of 6 to 7 heavy
cords with numerous fine spiral threads between the cords. Axial sculpture consisting of
2 or 3 coarsely knobbed varices, the knobs being formed where the spiral cords cross the

varices. In addition, there are 4 to 5 axially arranged and somewhat irregular knobbed

Five h,

— e
, ieee

<

Plate 118. Cymatium sarcostomum Reeve. Fig. 1. Pelican Shoals, Florida (2x). Fig. 2. Mauritius (2x). Fig.

3. One mile north of Frederiksted, St. Croix, Virgin Islands (1.6x). Fig. 4. Mauritius (1.6x). Fig. 5.

St. Croix, Virgin Islands (2.27x).

208 JOHNSONIA, No. 36 Cymatium

ridges between the varices. Operculum unknown. Periostracum generally deciduous,
light brown in color, thin and consisting of low, axial blades. Embryonic whorls unknown.

length width
60mm, 33.5 mm. Bermuda
56 37 Calapan, Mindoro, Philippine Islands

Types. The type specimens of C. sarcostomum Reeve, C. moritinctus Reeve, and C. rid-
leyi Smith are probably in the British Museum. The type locality for sarcostomum is the
Island of Ticao, Philippine Islands.

Remarks. This is a rare species in both the Western Atlantic and the Indo-Pacific re-
gions. Inthe Western Atlantic this species appears to be very closely related to C. carib-
hacum. The most significant differences separating these species are the fewer and more
prominent axial ridges of sarcostomum. In this species the axial ridges cross several of
the spiral cords while in C. caribbaeum the only evidence of the axial ridges is the align-
ment of the small knobs on the spiral cords.

Inthe Western Atlantic C. sarcostomum appears to occur in deeper water than it does
in the Indo-Pacific.

Range. WesvtERN ATLANTIC: From southern Florida and Bermuda through the West
Indies and south to Brasil. This last record is based upon C. ridleyi Smith as given in
the synonymy above.

Inpo-Paciric: From the Philippine Islands south through Indonesia and west to
Portuguese East Africa. Locard (1897) records this species (as moritinctum) from the
Cape Verde Islands, West Africa in the Eastern Atlantic.

Specimens examined. WESTERN ATLANTIC. FLoriIpA: off Palm Beach in 75 fathoms;
off Lake Worth in 70 fathoms; S. E. of Government Cut, Miami in 30 fathoms (all T.
McGinty); Pelican Shoals, off Key West (MCZ). Brermupa: (R. Merrill) [dredged
dead]. VirGin IsLanps: 1 mile N of Frederiksted, St. Croix (G. Usticke; R. Walker).
Lesser ANTILLES: off Sombrero Island in 45 fathoms (Blake, MCZ); Carenage, Trini-
dad (H. G. Kugler).

Inpo-Paciric. PHILIPPINE IsLANDs: Subic Bay, Zambales, Luzon; Zamboanga,
Mindoro (both MCZ). Moniucca Istanps: Morotai Island (MCZ). INp1an Ocean Is-
LANDS: Mauritius (MCZ); Porrucugsr East Arrica: Bazaruto Island, Bazaruto Bay
(J. IK. Howard).

Cymatium (Ranularia) pyrum rehderi Ve77i//
Plate 119, figs. 2-8

Cymatium rehderi Verrill 1950, Nautilus 63, p. 126, pl. 9, fig. la—b (25-40 fathoms off Dominica, British
West Indies [Lesser Antilles] ).

Description. Shell medium in size, reaching 92 mm. (about 84 inches) in length, solid,
rimately umbilicate and strongly sculptured. Color a light reddish brown, the varices
having alternating bands of brown and white. Six convex post-embryonic whorls. Spire
moderately extended and produced at an angle of 60°. Aperture subovate. Outer lip

Western Atlantic JOHNSONIA, No. 36 209

thickened and having seven rather coarse denticles. Inner lip consisting of a thickened
parietal area on which there are numerous, fine plicae which reach to the outer edge of
the thickened area. The spaces between the plicae are reddish brown. Siphonal canal
moderately broad and having a narrow aperture which is curved upward and slightly to
the left. Columella thickened and continued into the siphonal canal. Suture slightly in-
dented. Sculpture consisting of five or six varices, with five rather large, knobbed, axial
ridges between the varices and with numerous, fine axial threads. Spiral sculpture con-
sisting of about seven large cords interspaced with numerous threads. The crossing of the
axial and spiral threads creates a beaded appearance. Operculum, periostracum and em-
bryonic shell unknown.

Plate 119. Fig. 1. Cymatium pyrum Linné. Okinawa, Ryukyu Islands, Japan (1. 18x). Fig. 2. Cymatium pyrum
rehderi Verrill. Banes, Oriente, Cuba (slightly enlarged). Fig. 3. Cymatium pyrum rehderi Verrill, Dominica,
Lesser Antilles. Holotype, USNM no. 594095 (1.6x).

210 JOHNSONIA, No. 36 Cymatium

length width
59 mm. 30mm. Holotype of C. p. rehderi
92 46 Banes, Oriente, Cuba
65 33 Matanzas Bay, Cuba

Types. The holotype of Cymatium rehderi Verrill is in the United States National
Museum, no. 594095, from off Dominica, Lesser Antilles in 25-40 fathoms. The measure-
ments of the holotype given by Verrill in the original description were based on the photo-
graph which was enlarged and not the specimen itself. His original measurements gave
a length of 75 mm. and a width of 41 mm.

Remarks. Cymatium pyrum rehderi Verrill is very closely related to C. pyrum Linné
of the Indo-Pacific region. Actually the only significant differences between this sub-
species and the typical form are the smaller size of known specimens of C. p. rehderi, the
lighter color of its shell and the darker reddish brown of the parietal area. In C. pyrum
the parietal area is colored the same as the outer surface of the shell. We figure a speci-
men of C. pyrum (Plate 119, fig. 1) for comparison.

In the Western Atlantic C. p. rehderi is most easily confused with C. sarcostomum
Reeve from which it differs by having a more extended spire and having the plicae on
the parietal area extend to the outer edge of the thickened parietal lip. In addition, it
usually has a longer siphonal canal.

Range. This species appears to be restricted to the West Indies, occurring mainly in
fairly deep water.

Specimens examined. Cuna: Matanzas Bay and Varadero, Matanzas (both C.J. Finlay);
Habana, Habana; off Matanzas, Matanzas, Yara, station 8 (N. Lat. 23°8’; W. Long.
81°28’) in 200 fathoms (both Museo Poey); Banes, Oriente (C. J. Finlay); Guantanamo,
Oriente (USNM). Hispanro.a: Puerto Plata, Santo Domingo (MCZ). Lesser An-
TILLES: Dominica (USNM).

Subgenus Cymatriton, new subgenus

Shells rather small in size and generally uniformly colored a light ashen gray. Axial
sculpture consisting of several varices and large, knobbed, axial ridges. Spiral sculpture
consisting of large, rounded, cords interspaced with fine thread-like cords. Operculum
broadly oval, having a subcentral nucleus and sculptured with concentric growth lines.
Embryonic whorls convex, broad and minutely, axially striate.

Type species, Cymatium nicobaricum Réding.

Cymatium (Cymatriton) nicobaricum Réding
Plate 111, figs. 5-6; Plate 113, fig. 4; Plate 120, figs. 1-3

Tritonium nicobaricum Roding 1798, Museum Boltenianum, p. 126 [refers to Chemnitz 1780, in Martini,
Conchylien-Cabinet (1) 4, pl. 130, figs. 1246-1247].

Triton chlorostomum Lamarck 1822, Histoire Naturelle des Animaux sans Vertébres 7, p. 185 (1’Océan des
Antilles); Kiener 1842, Iconographie des Coquilles Vivantes 7, Triton, p. 19, pl. 12, fig. 2.

Triton pulchellus C. B. Adams 1850, Contributions to Conchology, no. 4, p. 60 (Jamaica).

Triton chlorostomum pumilio Mirch 1877, Malakozoologische Blatter 24, p. 29 (no locality given).

Western Atlantic JOHNSONIA, No. 36 211

Description. Shell medium to small in size, reaching 85 mm. (about 32 inches) in
length, solid, imperforate and strongly sculptured. Color a light ashen gray to rarely a
reddish brown. In addition, on most specimens examined, there are flecks of reddish
brown, usually appearing on the coarse spiral threads. Interior of aperture a bright orange,
the lamellae and palatal denticles white. Occasional specimens have the aperture entirely
white. Post-embryonic whorls 7 and strongly convex. Spire extended and produced at
an angle of about 45°. Aperture subcircular, with the outer lip greatly thickened when
a varix is produced. Parietal lip rather narrow, heavily glazed and with numerous, some-
what irregular, low lamellae. Inner margin of the outer lip with 7 strong denticles which
extend well back into the aperture and which may be single or divided. Columella
arched inward, with its base continuing into the parietal margin of the siphonal canal.
Siphonal canal narrow, moderately long and generally curved upward. Suture slightly
indented. Sculpture consisting of 6 strong spiral cords, which are nodulose and are inter-
spaced with fine thread-like cords. Axial sculpture consisting of 5 to 8 rounded varices.
There are 3 to 5 larger knobs between each pair of varices. These knobs are variable in
size, both as to length and height. Many of them may extend over three of the heavy
spiral cords. Periostracum reddish brown, very thin, produced in fringed axial blades
and usually deciduous. Operculum broadly oval, with a subcentral nucleus and sculp-
tured with concentric growth lines. Embryonic whorls 6, strongly convex, broad, amber
in color and with fine axial striae.

length width
85 mm. 49mm. Marquesas Islands, Polynesia
78 43 Oshima, Osumi, Ryukyu Islands, Japan
71 43 Bermuda
Lees 29 Punta de los Colorados, Cienfuegos, Cuba
eat 34 Long Reef, off Elliott Key, Florida

Types. The location of the type specimen of 7°. nicobaricum Roding is unknown. The
type figure is in the Conchylien-Cabinet (1) 4, pl. 180, figs. 1246-1247. The holotype
of 7. chlorostomum Lamarck is probably in the Paris Museum. The holotype of 7". pul-
chellus C. B. Adams is in the Museum of Comparative Zoology, no. 186135. We here
restrict the type locality of C. nicobaricum to Jamaica.

Remarks. Like others in this family, this is a variable and widely dispersed species,
occurring in both the Western Atlantic and the Indo-Pacific regions. We have been un-
able to separate the Indo-Pacific specimens from those of the Western Atlantic. The
greatest variability is expressed in the number and size of the knobs. Some specimens
have but a few large and rather irregular knobs while others have numerous, small and
quite regular ones.

This species is not closely related to any other in the Western Atlantic. It superficially
resembles both C. pyrum rehderi and C. pileare. However, the extended spire and small
and nearly circular aperture of nicobaricum readily differentiates it from rehderi. From
C. pileare it differs in having a nearly circular rather than oval aperture, a more twisted
siphonal canal and a much coarser, knobbed sculpture.

The embryonic whorls in this species are quite different from those of most other spe-
cies in the genus Cymatium in the Western Atlantic. hey are proportionately much
wider, stouter and far more conical in shape than other embryonic shells in this genus.

JOHNSONIA, No. 36 Cymatium

io
m

Range. WersvteERN ATLANTIC: From Palm Beach County, Florida, south to Tortugas;
Bermuda and the West Indies, and from Mexico south at least to Honduras. It has been
reported from Bahia, Brasil by Lange de Morretes (1949, p. 92).

INpo-Paciric: Hawaiian and Marquesas Islands, west to southern Japan, south to
New Caledonia and west, through the East Indies and the Indian Ocean Islands to
Mauritius. It has been reported from New South Wales, Australia according to D. F.
McMichael (in litt.). There are several published records of this species from Madeira
and the Canary Islands in the Eastern Atlantic.

Specimens examined. WESTERN ATLANTIC. FLortpa: Jupiter Inlet; Riviera Beach,
Palm Beach County (both FSM); Lake Worth, Boynton (FSM; T. McGinty; G. Kline;
J. Schwengel); Boca Raton Inlet (J. A. Flowers); Bear Cut, Biscayne Key (H. and K.
Johnstone): Long Reef, off Elliott Key (G. Voss); Molasses Reef, off Key Largo (R.

Figs. 1-2. Nuevitas, Camagiiey, Cuba (2x). Fig. 3. Barbados,

2)

Plate 120. Cymatium nicobaricum Rédin

go,
5
2
oO.

‘

Lesser Antilles (embryonic whorls, 1

Western Atlantic JOHNSONIA, No. 36 213

Work): Sombrero Key (MCZ); Looe Reef, off Big Pine Key (MCZ); Sand Key, Key
West (FSM): Pelican Shoals, Key West (R. Work); Garden Key, Dry Tortugas
(FSM). Bermupa: (ANSP; MCZ). Banama Istanps: West End Point, Grand Ba-
hama Island (MCZ); Hope Town, Great Abaco (R. Robertson); Sweetings Village,
Great Abaco; Alice Town, North Bimini, Bimini Islands (both MCZ); North Cat Cay
and Gun Cay, Bimini Islands; Morgans Bluff, Andros; Clifton Bluff, New Providence
(all T. McGinty); Dicks Point, New Providence (MCZ); Harbour Island, Eleuthera
(G. Kline); Little San Salvador Island; Northeast Point, 4 miles E of Arthurs Town,
Cat Island; Clarence Town, Long Island; Abrahams Bay, Mariguana Island; Matthew
Town, Great Inagua (all MCZ). Cupa: Habana, Habana (ANSP); Matanzas Bay,
Matanzas (C. J. Finlay); Varadero, Cardenas (C. G. Aguayo); Playa Comancho, Car-
denas (C. J. Finlay); Cayo Santa Maria, off Punta Alegre, Camagiiey (R. Humes):
Nuevitas, Camagiiey (C. J. Finlay); Castillo de Jagua, Cienfuegos; Punta de los Col-
orados, Cienfuegos (both MCZ); Rente, Santiago de Cuba(C. G. Aguayo); Fish Point,
Guantanamo Bay (MCZ). Hispanrtota: Miragoane, Haiti (W. J. Eyerdam); Monte-
cristi; Puerto Plata; Puerto Sostia; Santa Barbara de Samana, all Santo Domingo (all
MCZ). Jamaica: Port Antonio (MCZ); Whitehouse (J. K. Howard). Purrro Rico:
Mona Island (A. Phares); San Juan; San Geronimo; Camuy; Rincé6n; Bahia de An-
asco; Punta Algarrobo; Mayagiiez; Punta Cuchara; near Maunabo (all G. Warmke);
Ponce (MCZ). VirGin Istanps: The Baths, Virgin Gorda; Guana Island, Tortola
(both M. W. Dewey); St. John (MCZ); St. Thomas (MCZ; ANSP); 1 mile N of
Frederiksted, St. Croix (G. Usticke). Lesser ANTILLES: Barbados; Speyside, Tobago;
Carenage, Trinidad (all MCZ). CaripBpean IsLANpDs: Georgetown, Grand Cayman, Cay-
man Islands (C.G. Aguayo); Roatan Island, Bay Islands (ANSP). Mexico: Tuxpan,
Veracruz; Veracruz, Veracruz (both T. Pulley); Cabo Catoche, Yucatan (MCZ). Hown-
puRAS: Balfate (FSM).

Inpo-Paciric. HawatiaAN Istanps: Hilo, Hawaii (MCZ; J. Schwengel); Kihui,
Maui (W. Old); Waikiki, Oahu; off Koko Head, Oahu (both MCZ); Fort Kameha-
meha, Oahu (J. Schwengel); Midway Island (MCZ). Line Isuanps: Palmyra Island
(MCZ). Marquesas Istanps: (MCZ). Socrety IsLANDs: Punaauia, Tahiti; Vaitape,
Bora Bora (both R. Robertson). PHorNnrx IsLanps: Canton Island (MCZ). Samoan
IsLtanps: Nuuuli, Tutuila (MCZ). Exviice Isuanps: Funafuti (MCZ). Fis1 IsLanps:
Bega Island (J. Schwengel); Tokoh, 3 miles SW of Levuka, Ovalau (MCZ). GiLBerr
Istanps: Apiang (MCZ). Marswauui Istanps: Ebon Island (MCZ); Rujoru Island
and Iqurin Island, Eniwetok Atoll; Bikini Island, Bikini Atoll; Naen Island; Ronge-
lap Atoll (all USNM). Marranas Istanps: Agana Bay, Guam; Tinian Island; ‘Tumon
Bay, Guam (all MCZ); Saipan (W. Old). Caro.ine Isuanps: N of Garokottan Island;
Garakayo Island; Ngargersiul Island; East Babelthuap Island, all Palau Islands (all
ANSP). New Careponia: Isle of Pines (MCZ). New Guinea: Gusika, 13 miles N
of Finschhafen, Australian New Guinea; Jamna Island, Dutch New Guinea (both
MCZ). Abroeki, Aoeri Island, Geelvink Bay, Dutch New Guinea (ANSP). Aus?rRa.ia:
Lizard Island, north Queensland (E. Grigg). Gloucester Island, Whitsunday Group,
Queensland (W. Old). Japan: Tokyo Bay (MCZ). Ryukyu IsLtanps: Oshima, Osumi
(MCZ). Puinipeixe Isutanps: Guiuan, Samar Island: Lubang Island; Calapan, Min-
doro; Siasi Island, Tapul Group, Sulu Islands (all MCZ). Monucca IsLtanps: Bouro Is-
land; Amboyna; Tengah Island, Bouro Island; Dagasoeli Island, North Loloda Island;

214 JOHNSONIA, No. 36 Cymatium

Morotai Island (all MCZ). INp1AN OcEaN IsLanps: Mauritius (MCZ; J. Schwengel).

Subgenus Septa Perry

Septa Perry 1810, Arcana, pl. 2, fig. 2; Perry 1811, Conchology, London, pl. 14, fig. 2 (type species,
Septa scarlatina Perry [ = Murex rubeculus Linné], monotypic).
Lampusia Schumacher 1817, Essai Nouveau Systéme, p. 350 (type species, Murex pilearis Linné, subsequent

designation, Herrmannsen 1847).
Simpulum Morch 1852, Catalogus Conchyliorum Comes de Yoldi, p. 108 (type species, Murex rubeculum

Linné, here selected); non Simpulum Fabricius 1823.
Simplum Stoliczka 1867, Paleontologia Indica (5) 2, p. 131 (error for Simpulum Morch).

Type species, Septa scarlatina Perry [= Murew rubeculus Linné], monotypic.

Shells medium to small in size, exceptional shells of C. pileare Linné reaching 188 mm.
(about 54 inches) in length; attenuate, imperforate and generally solid in structure.
Color white buff or gray usually with spiral bands of brown or red-brown. Sculpture
very variable among the several species, but generally with well-developed varices and
strong spiral cords or ribs. Many species have fairly large knobs between the varices and
these can be in axial arrangement. Aperture generally with well-developed denticles on
the outer lip and many, usually fine plicae on the parietal wall. Siphonal canal short
and usually curved upward.

Cymatium (Septa) rubeculum occidentale, new subspecies
Plate 110, fig. 3; Plate 113, fig. 5; Plate 121, fests

Triton rubeculum occidentale ‘Mirch’ Tryon 1881, Manual of Conchology (1) 3, p. 12 [nomen nudum].

Description. Shell rather small, reaching 32 mm. (about 14 inches) in length, rather
solid, imperforate and strongly sculptured. Color light brownish yellow with 1 to 2 spiral
bands of white. Post-embryonic whorls 5. Spire moderately extended and produced at
an angle of about 55°. Aperture subelliptical, the outer lip greatly thickened when a
varix is produced. Outer lip with 8 denticles. Inner lip with 14 or 15 rather coarse lam-
ellae. Siphonal canal rather narrow, short and curved upward. Columella nearly straight.
Suture slightly indented. Sculpture consisting of 7 or 8 beaded cords of nearly equal
strength. Axial sculpture consisting of 8 to 5 ribbed varices. Between the varices there
are 3 to 5 weak axial ridges. In addition, there are close-set axial threads which, cross-
ing the spiral cords, produce the beaded effect and give the shell a reticulated appearance.
Periostracum light straw-yellow in color and consisting of numerous close-set, low, axial,
fringed blades. Operculum unknown. Embryonic shell very small for the size of the
adult and consisting of 8 whorls which are light brown in color.

length width
32 mm. 17.5mm. Holotype, St. Thomas, Virgin Islands
25 16 Paratype, Carboneras, Matanzas, Cuba

Types. The holotype is in the Academy of Natural Sciences Philadelphia, no. 36874,
from St. Thomas, Virgin Islands, R. Swift, collector. Paratypes from Carboneras, 14
miles west of Rio Camarioca, Matanzas, Cuba, collected by C. J. Finlay are in the collec-
tions of C. J. Finlay, T. McGinty and the Museum of Comparative Zoology, no. 202269.

Western Atlantic JOHNSONIA, No. 36 215

Remarks. We are describing C. rubeculum occidentale as a new subspecies though this
has long been known under the name occidentale Mérch. However, Mérch never de-
scribed this subspecies. He used the term ‘‘occidentale’” four times under the genus
Triton in his paper on the ‘‘Synopsis molluscorum marinorum Indiarum occidentalium’’
(1877, Malakozoologische Blatter 24, pp. 14-52). His use of the word ‘‘occidentale”’
was to indicate that he was considering only the specimens from the West Indies and
not the Orient. ‘Tryon made the original error in the Manual of Conchology (loc. cit.)
when he stated that ‘‘Mérch made a variety, occidentale, but it has no distinctive charac-
ters.”’ So far as we can determine no subsequent author referring to this form has ever
described or figured it.

The subspecies occidentale differs from the typical form by being smaller, having a finer
sculpture and being more somberly colored. Typical rubeculum of the Indo-Pacific region
is usually an intense brick-red or orange-brown, while in occidentale the color is usually
a dull brownish red toa light brown. In the West Indies this subspecies could be easily
confused with small specimens of C. pileare, but they may be separated by the obtuse
point of the spire and beaded sculpture of occidentale.

It would appear that the embryonic shell breaks off very easily in this subspecies. We
have seen a few specimens which were collected alive but even these lack the embryonic
whorls. Only a single specimen with the embryonic shell still attached has been seen.

Range. From southern Florida and probably all of the West Indies west to Central
America.

Plate 121. Cymatium rubeculum occidentale Clench and Turner. Fig. 1. St. Thomas, Virgin Islands (Holotype,
2.8x). Fig. 2. Pigeon Cays, Andros Island, Bahama Islands (3x). Fig. 3. Off Lantana, Florida in 10 fath-

oms (embryonic whorls, 10x).

216 JOHNSONIA, No. 36 Cymatium

Specimens examined. Fitoripa: Palm Beach in 25 and 40 fathoms (both T. McGinty) ;
Ocean Ridge and off Lantana in 10 fathoms, Palm Beach County (both FSM). Banama
IsLANps: Pigeon Cays, Andros (T. McGinty). Cusa: Carboneras, 14 miles W of Rio
Camarioca, Matanzas (C.J. Finlay; T. McGinty; MCZ). Viren Istanps: St. Thomas
(ANSP). Mexico: Cabo Catoche, Yucatan (MCZ); Isla Mujeres, Yucatan (Museo
Poey).

Cymatium (Septa) pileare Linné
Plate 112, figs. 1-2; Plate 118, fig. 7; Plate 122, figs. 1-8; Plate 123

Murex pileare Linné 1758, Systema Naturae, ed. 10, p. 749 [refers to Gualtierius 1742, pl. 49, fig. G] (M.
Mediterraneo).

Triton pileare Linné, Lamarck 1816, Tableau Encyclopédique et Méthodique 3, pl. 415, fig. 4a—b; Liste,
p. 4; Lamarck 1822, Histoire Naturelle des Animaux sans Vertébres 7, p. 182 (1’Océan des Antilles).

Triton aquatilis Reeve 1844, Conchologia Iconica 2, Triton, pl. 7, fig. 24 (Ticao, Philippine Islands).

Triton martinianum d’Orbigny 1847 [in] Sagra, Histoire de l’Ile de Cuba, Mollusques 2, p. 162 [new name
for Triton pileare Lamarck 1822, non Murex | Triton] pileare Linné 1758].

Litiopa effusa C. B. Adams 1850, Contributions to Conchology, no. 5, p. 71 (Jamaica); Turner 1956, Occa-
sional Papers On Mollusks 2, p. 136, pl. 21, fig. 3 [is the veliger stage].

Triton intermedius Pease 1869, American Journal of Conchology 5, p. 74 (Oahu, Hawaiian Islands).

Triton martinianum latior Morch 1877, Malakozoologische Blatter 24, p. 29 [nomen nudum].

Triton veliei Calkins 1878, Records and Proceedings Davenport Academy of Natural Sciences 2, p. 235,
pl. 8, figs. 1-2 (southern Florida).

Dissentoma prima Pilsbry 1945, Nautilus 59, p. 59, text fig. 1 (off Singers ane near North Inlet, Lake
Worth, Palm Beach, Florida); Pilsbry 1949, Nautilus 62, p. 142.

Description. Shell moderately large, reaching 138 mm. (about 54 inches) in length,
solid, imperforate and strongly sculptured. Color, in specimens denuded of their perios-
tracum, a grayish brown to golden brown, generally banded with alternating light and
dark bands, particularly on the varices. Within the aperture the color on the outer lip
is reddish-brown with 12 to 14 whitish plicae which are often paired. Parietal wall a dark
chocolate in color, with numerous fine, irregular, white lamellae. Post-embryonic whorls
seven and strongly convex. Spire moderately extended and produced at an angle of 45°.
Aperture elliptical with the outer lip greatly thickened when a varix is produced. Pari-
etal lip glazed and with numerous irregular lamellae. Siphonal canal short and generally
curved upward. Columella arched inwardly with its base continuing into the parietal
margin of the siphonal canal. Suture slightly indented. Sculpture consisting of numerous
spiral and generally nodulose cords of unequal strength. Axial sculpture consisting of
3 to 5 strongly knobbed varices. Periostracum generally a light golden brown in color,
roughened with numerous periostracal ‘hairs’ and axial blades. Operculum unguiculate
with a marginal nucleus and sculptured with numerous concentric growth ridges. Em-
bryonic whorls four, slightly convex, amber in color and with very fine, axial striae.

length width
138 mm. 65mm. Philippine Islands
115 56 Rebecca Shoals, 45 miles W of Key West, Florida
111 OAs) Banes, Cuba
100 AT Mauritius
96 47.5 Castle Harbour, Bermuda
94. 44 Boca Raton, Florida

84.5 42.5 Biscayne Bay, Florida

ae ee

Western Atlantic JOHNSONIA, No. 36 2

Types. The only reference given by Linné for pileare was to Gualtierius, pl. 49, fig.
G and this is the type figure. The locality ‘“‘M. Mediterraneo’ given by Linné is in error.
This species does not occur in the Mediterranean. The lectotype of Litiopa effusa C.B.
Adams is in the Museum of Comparative Zoology, no. 186589 from Jamaica. The lec-
totype of 7". intermedius Pease is also in the Museum of Comparative Zoology, no.
191331, from Oahu, Hawaiian Islands. The holotype of Dissentoma prima Pilsbry is in
the Academy of Natural Sciences Philadelphia, no. 181369. The location of the holo-
type of 7° veliei Calkins is unknown to us, but idiotypes from Key West, Florida are in
the Museum of Comparative Zoology, no. 150085.

Plate 122. Cymatium pileare Linné. Fig. 1. Apiang Island, Gilbert Islands (2x). Fig. 2. Oahu, Hawaiian
Islands (lectoty pe of Triton intermedius Pease | =C. pileare Linné] MCZ 191331, about 2x). Fig. 3. Venetian

Causeway, Biscayne Bay, Florida (1.35x).

218 JOHNSONIA, No. 36 Cymatium

As Linné’s type locality, as stated above, was in error we take that given by Lamarck
as ‘l’Océan des Antilles’ and restrict it to Jamaica, the locality given by Lister who
was Lamarck’s first reference.

Remarks. This is an exceedingly far-ranging species occurring as it does in the tropical
portion of the Western Atlantic and throughout most of the Indo-Pacific. There have
been several attempts to separate specimens of the Atlantic from those of the Indo-
Pacific, beginning with d’Orbigny who created a new name for Lamarck’s use of pileare
Linné. However, we have been unable to separate specimens from these two very dis-
tant areas on any shell characters.

Individual specimens from one locality may show a great deal of variation, but there
does not appear to be any geographic significance to these variations.

Range. WrsTERN ATLANTIC: From Jupiter Inlet, Florida south to Tortugas; off
Port Aransas, Texas in deep water (32 fathoms); Bermuda and the West Indies from
the Bahamas, and from Veracruz, Mexico south to Bahia, Brasil.

Inpo-Paciric: The Hawaiian Islands, west to the Ryukyu Islands, Japan and south
through the East Indies and the Indian Ocean to East Africa.

Specimens examined. WrsTERN ATLANTIC. FLoripa: Jupiter Inlet (FSM); Boynton
Inlet, Lake Worth (D. Moore; FSM; T. McGinty): 5 miles S of Delray Beach (FSM);
Boca Raton Inlet (EK. V. Flowers; FSM: T. McGinty); Pompano (MCZ); Biscayne
Bay (R. Humes): off Key Largo (R. Merrill); off Sombrero Light in 85 fathoms (T.

= a a

Plate 123. Cymatium pileare Linné. Arthurs Town, Cat Island, Bahama Islands (embryonic whorls, 36x).

Western Atlantic JOHNSONIA, No. 36 219

McGinty); Bahia Honda Key (D. and N. Schmidt); Pelican Shoal, off Boca Chica
Key; Key West; Sand Key, off Key West (all MCZ); off Rebecca Shoals, about 45
miles W of Key West in 19 fathoms (H. and K. Johnstone); Dry Tortugas (MCZ).
Texas: SE of Port Aransas in 32 fathoms (C. L. Branch). Bermupa: Castle Harbour
(dredged fossil, H. B. Moore). BaHAMaA IsLanps: Eight Mile Rock, Grand Bahama
(MCZ); Cooper Jacks Cays, Great Abaco (R. Robertson); Alice Town, North Bimini
(MCZ); Gun Cay, Bimini (T. McGinty); Middle Bight, Andros; Dicks Point, Nassau,
New Providence (both G. Kline); Savannah Sound, Eleuthera; Orange Creek, near
Arthurs Town, Cat Island; Clarence ‘Town, Long Island; Watling Island; Fortune
Island; Cay Sal, Cay Sal Bank (all MCZ). Cupa: Puerto Esperanza, Pinar del Rio;
Habana, Habana; Camarioca, Matanzas; Matanzas, Matanzas (all Museo Poey):; Vara-
dero, Matanzas (J. Finlay); Caibarién, Las Villas (MCZ); Cayo Maja Figuro, Punta
Alegre, Camagiiey (R. Humes); Banes, Oriente (J. Finlay); Levisa Bay, Mayari, Ori-
ente (A. Quifiones); Maisi, Oriente (Museo Poey); Cuesco Beach, Guantdanamo Naval
Base (MCZ); Nispero, Santiago, Oriente (Museo Poey); Caleton de Don Bruno, Cien-
fuegos (MCZ). Jamaica: Port Antonio(MCZ; ANSP); Whitehouse (J. K. Howard).
HispaNIo_a: Jérémie and Miragoane, Haiti; Montecristi, Puerto Plata, and Puerto
Sostiia, Santo Domingo (all MCZ). Purrro Rico: Mona Island (MCZ): San Juan
(D.Thomas; R.Work); Arecibo (G. Warmke); Bahia de Aguadilla (A. Phares); Rin-
con; Bahiade Afiasco; Mayagiiez; Punta Guanajibo; Guanica (all G. Warmke; MCZ):
Ponce (MCZ); near Maunabo (G. Warmke). VirGin Istanps: Frederiksted, St. Croix
(G. Usticke; MCZ); St. John; St. Thomas (both MCZ); Guana Island, Tortola;
The Baths, Virgin Gorda (both M. W. Dewey). LrEssErR ANTILLES: St. Kitts; Guade-
loupe (both ANSP); St. Lucia; Barbados (both MCZ); Buccoo Reef, Tobago (MCZ;
K. Anderson); Chaguaramas, Trinidad (H.G. Kugler); Grenada (K. Anderson). Carrp-
BEAN IsLanps: Southwest Point, Grand Cayman, Cayman Islands (ANSP). Mexico:
Tuxpan, Veracruz; Cabo Catoche, Yucatan; Isla Mujeres, Yucatan (all MCZ). Hon-
puRAS: Puerto Cortez (MCZ); Belfate (FSM). Panama: 2 miles off Col6n; Brujas
Point, Canal Zone (both W.G.Clark). Durcu Gurana: Corentyne River (dead, H.G.
Kugler); Brasii: Itaparica, Ilha de Itaparica, Bahia (H.S. Lopes).

INpo-Paciric. Hawaan Istanps: Fort Kamehameha, Oahu (J.Schwengel); Pearl
Harbor, Oahu (V.D.Spicer). Lint Isuanps: Kingman Reef, 33 miles NW of Palmyra
Island (J. Schwengel). Sociery Istanps: Arue, Tahiti; Vaitape, Bora Bora (both R.
Robertson). GitBert IsLtanps: Apiang (MCZ). CaroLine Istanps: Garakayo Island,
Palau Islands; Babelthuap, Palau Islands (both ANSP). Marianas Isutanps: Saipan
(W.Old). Fis1 Istanps: Bega Island (J.Schwengel). Japan: Ryukyu Islands (MCZ).
PHILIPPINE IsLANDS: Subic Bay, Zambales, Luzon; Calapan, Mindoro; Tagbac, Lu-
bang; Araceli, Dumaran Island, Palawan; Davao, Mindanao; Zamboanga, Mindanao
(all MCZ). New Guinea: Island 5 miles NW of Rani Island, Schouten Islands; Ab-
roeki, Aoeri Island, Geelvink Bay; Samberbaba, Japen Island, all Dutch New Guinea
(all ANSP); Jamna Island, Dutch New Guinea (MCZ); Madang Island, Finschafen,
Australian New Guinea (E. Grigg). AusrraLtia: Augustus Island, northern Western
Australia (MCZ): Low Wooded Isle and Portland Roads, both northern Queensland
(both E. Grigg). INpones1a: Toelehoe, Amboina (MCZ); Woda Island, Halmahera,
Molucea Islands; Moratai Islands; St. Nicholas Bay, Bali Island, Greater Sunda Islands
(all MCZ). Inp1an Ocean IsLanps: 12 miles N of Trincomalee, Ceylon (W. Old);

220 JOHNSONIA, No. 36 Cymatium

Mauritius; Zanzibar (both MCZ):; Nosy Bé, Madagascar (A. Humes). Pakistan: Mak-
ran Coast (MCZ). ApEN: (MCZ). Kenya: Mombassa (MCZ); Kilifi (E. Grigg).
PorrucuesE East Arrica: Bazaruto Island, Bazaruto Bay (J. K. Howard).

Cymatium (Septa) krebsii Maérch
Plate 112, figs. 3-4; Plate 124, figs. 1-4

Triton krebsii Mirch 1877, Malakozoologische Blatter 24, p. 30 (St. Thomas and St. Croix [Virgin Islands] );
Kobelt 1878, Conchylien Cabinet (2) 3, pt. 2, p. 265, pl. 70, figs. 3-4.

Cymatium (Lampusia) krebsii Mérch. Rehder and Abbott 1951, Revista de la Sociedad Malacolégica 8, pl. 8,
fig. 6.

Lampusia? pharcida Dall 1889, Bulletin Museum Comparative Zoology 18, p. 227, pl. 36, fig. 2 (Blake,
station 293, N. Lat. 13°14/23/"; W.Long. 59°39/10”, off Barbados in 81 fathoms).

Description. Shell medium in size, reaching 73 mm. (about 3 inches) in length, solid,
imperforate and strongly sculptured. Color entirely white or white with spiral bands of
yellowish brown in the grooves between the cords. Interior of the aperture white. Post-
embryonic whorls 7 and strongly convex. Spire extended and produced at an angle of
about 37°. Aperture nearly vertical and narrowly elliptical. Outer lip greatly thickened
when a varix is produced. Parietal lip rather narrow and heavily glazed. On the outer
lip there are 6 or 7 strong denticles which extend well into the aperture. On the inner
lip, at the upper end there is a strong lamella which forms the margin of the anal canal.
On the columella and well within the aperture there are two large and somewhat irreg-
ular lamellae. In addition, between the anal lamella and the base of the siphonal canal
there are numerous and very fine thread-like lamellae. Columella slightly arched in-
wardly, with its base continuing into the parietal margin of the siphonal canal. Siphonal
canal narrow, rather short and curved upward. Suture slightly indented. Sculpture con-
sisting of from 5 to 7 strong, nodulose varices. Between the varices there are 5 or 6
somewhat irregular and strongly nodulose, axial ridges. These are crossed by 6 or 7
heavy spiral cords. Between these cords there are three subequal but rather fine thread-
like cords. Periostracum thin, deciduous, roughened and golden-brown in color. Oper-
culum unguiculate with a marginal nucleus and numerous concentric growth lines.
Embryonic whorls 34 to 4, smooth, straw-yellow in color, narrow, extended and with
microscopic axial striae.

length width
73mm. 37mm. off New Ground Shoals, Dry Tortugas, Florida
58.5 29 Punta Guanajibo, Puerto Rico

Types. The holotype of C. krebsii Mérch is probably in the Universitetets Zoologiske
Museum, K6ébenhavn, Denmark. There are two paratypes from the Swift Collection in
the Academy of Natural Sciences Philadelphia, no. 42605. The holotype of C. pharcidum
Dall is in the United States National Museum, no. 94887. The type locality is here re-
stricted to St. Thomas, Virgin Islands, the locality from which Swift’s material came.

Remarks. Cymatium krebsii Morch is a very rare species to judge by the few specimens
in museum collections. It differs from C. nicobaricum in the shape of the aperture and
the lack of color in the aperture, though in general sculptural characters it appears to be
quite close to that species. From C. pileare Linné it differs in general shape and in the

i ae) ae

Western Atlantic JOHNSONIA, No. 36 99]

lack of color in the aperture. In addition, it is far more nodulose. From both C. nico-
baricum and C. pileare it differs by having two rather large columellar lamellae. See also
Remarks under C. gemmatum Reeve.

Cymatium krebsii is apparently related to C. corrugatum Lamarck of the Eastern At-
lantic, both species agreeing in the approximate shape of the more or less vertical and
narrow, elliptical aperture. In both species the aperture is white and both have the large
columellar lamellae. C. Akrebsu differs from corrugatum in being somewhat smaller and
in being far more nodulose.

Range. From Palm Beach County, Florida to Dry Tortugas and south through the
West Indies.

Records. Fiorina: off Delray Beach in 65 and 80 fathoms (FSM; T. McGinty); off
Palm Beach in 25 to 75 fathoms (T.McGinty; J. Schwengel; ANSP); Nellies Point,
Fish Haven, Palm Beach County (ANSP); North Inlet, Lake Worth (T. McGinty);
SE of Miami in 30 fathoms (T. McGinty); Little Duck Key (J. Schwengel); off Som-
brero Light, E of Marathon in 37 fathoms (T. McGinty); Bahia Honda Key; off Key

Plate 124. Cymatium krebsii Mérch. Fig. 1. Off Sombrero Key, Florida (about 1.4x). Fig. 2. St. Thomas,
Virgin Islands (paratype, 1.32x). Fig. 3. Off Barbados, Lesser Antilles (Holotype of C. pharcidum Dall [=
C. krebsii Mérch], 5x). Fig. 4. Lake Worth, Palm Beach Co., Florida (showing embryonic whorls, 8x).

2929 JOHNSONIA, No. 36 Cymatium

West in 18 fathoms (both H. and K. Johnstone); off Sand Key Light, Key West in 70
fathoms (‘T. McGinty); off New Ground Shoals, Dry Tortugas (USNM); off Tortugas
in 18-20 fathoms (T. McGinty); off Destin in 14 fathoms (ANSP: T. McGinty); off
Panama City in 21 fathoms (T. McGinty). Purrro Rico: Punta Guanajibo (A. Phares).
Vircin Isuanps: St. Thomas (ANSP). Lesser ANTILLES: off Scarborough, Tobago
in 36 fathoms (MCZ). Venezue.a: Cubagua Island (MCZ).

Cymatium (Septa) gemmatum Peeve
Plate 110, fig. 2; Plate 113, fig. 6; Plate 125, figs. 1-2

Triton gracilis of authors, not of Reeve 1844.
Triton gemmatus Reeve 1844, Conchologia Iconica 2, Triton, pl. 15, fig. 60 a-b (Island of Ticao, Philip-

pine Islands).
Triton mundum Gould 1849, Proceedings Boston Society Natural History 3, p. 143 (Tutuila, Samoan Is-

lands).

Description. Shell small, reaching 43:mm. (about 1# inches) in length, rather solid,
imperforate and strongly sculptured. Color yellowish brown, with patches of reddish
brown on the varices. Interior of the aperture white. Post-embryonic whorls six and
convex. Spire extended and produced at an angle of about 50°. Aperture elliptical.
Outer lip thickened, the inner edge with 12 denticles which are grouped in pairs. Upper
portion of the parietal wall with a large tooth formed to create the anal canal. Central
portion of the parietal wall with very weak lamellae. At the base of the aperture the
parietal lamellae increase in size and then diminish along the siphonal canal. Siphonal
canal long, narrow, almost closed and curved slightly upward. Suture slightly indented.
Axial sculpture consisting of 2 or 8 varices. Between the varices there are 3 or 4 rather
weak axial ridges; in addition, there are numerous very fine, axial threads. Spiral sculp-
ture consisting of numerous moderately heavy cords which are interspaced with finer
cords. Embryonic whorls 5, yellowish amber in color, rather narrow and glass-like. Oper-
culum unguiculate with a marginal nucleus at the base and with concentric growth lines.
Periostracum thin, straw-yellow in color, smooth over most of the shell but producing
fringed blades on the varices and axial ribs.

length width
43.5 mm. 21mm. Oshima, Osumi, Japan
36 18 Bazaruto Bay, Portuguese East Africa
34 14 Cays off Cardenas, Matanzas, Cuba
rt S) 13.5 Puerto Plata, Santo Domingo

Types. The types of C. gemmatum Reeve are probably in the British Museum; the
type locality is the Island of 'Ticao [off Masbate], Philippine Islands. The location of the
holotype of C. mundum Gould is unknown. An idiotype is in the Museum of Com-
parative Zoology.

Remarks. This species has long been known under the name of gracilis Reeve, but it
certainly does not fit the description and figure of that species. Reeve’s figure gives no
indication of the enlarged columellar plicae and the large parietal lamella which aids in
forming the clearly defined anal canal. Both of these characters are exhibited in the fig-
ure of gemmatum.

Western Atlantic JOHNSONIA, No. 36 293

This is a rare species; it has been taken from low water to depths of 117 fathoms.

In relationship it appears to be nearest to C. hrebsii Moérch. It differs from that species
by having a smaller aperture and by having the columellar plicae proportionately smaller.
Adult specimens of C. gemmatum are very much smaller, being only about half the size
of an adult C. krebsu. However, it is often difficult to differentiate the young of krebsii
from adult gemmatum.

The range given below is only an indication of the range of this species. It probably
occurs throughout all of the tropical portions of both the Indo-Pacific and the Western
Atlantic.

Range. WESTERN ATLANTIC: Southern Florida from Palm Beach County to the
lower Florida Keys and the West Indies, south to Trinidad.

Inpo-Paciric: The Hawaiian Islands, west to southern Japan, south to northern
Australia and west to Portuguese East Africa.

Specimens evamined. WESTERN ATLANTIC. FLORIDA: Boynton Beach (T. McGinty):
off Boynton, Palm Beach County (from stomach of a moray in 10 fathoms); 2 miles S
of Boynton in 10 fathoms (both FSM); off Lantana in 25 fathoms; off Palm Beach In-
let in 25 and 40 fathoms; Boca Raton (all T. McGinty); off Hillsborough Light, Brevard
County in 20 fathoms (FSM); off Miami in 27 fathoms (‘T. McGinty); off The Elbow,
Key Largo in 66 fathoms; 5 miles off Carysfort Light, Key Largo in 96 and 117 fath-
oms (all MCZ). Bermupa: St. George (T. McGinty). Banama ISLANDs: Channel Cay,
Great Abaco (R. Robertson); Nassau and Clifton Bluff, New Providence; North Bimini

and Gun Cay, Bimini Islands (all T. McGinty). Cupa: Arenas de la Chorrera, Habana

Plate 125. Cymatium gemmatum Reeve. Fig. 1. Cardenas Keys, Matanzas, Cuba (2.9x). Fig. 2. North Bim-
ini Island, Bimini Islands, Bahamas (8x).

224, JOHNSONIA, No. 36 Cymatium

(MCZ): Cays off Cardenas, Matanzas: Varadero Beach, Matanzas (both C. J. Finlay).
HispanioLta: Cap Haitien, Haiti (T. McGinty): Caye Carenge, Boucassin, Haiti (R.
Work); Puerto Plata, Santo Domingo (MCZ). Purrro Rico: Arecibo; Punta Algar-
robo (both U.P.); Rincén (G.Warmke). Viren Istanps: St. Croix (FSM). Lesser
ANTILLES: off Barbados, Blake, in 50 and 100 fathoms; 2 miles S of Fort George,
Scarborough, Tobago in 36 fathoms (both MCZ).

Inpo-Paciric. HawalltaANn IsLanps: off Koko Head, Oahu(MCZ). Samoan IsLanps:
Vaoto, Vailele Bay, Upolu Island (ANSP). Hownanp Istanp (MCZ). Ginperr Is-
LANps: Apiang (MCZ). Caronine IstaAnps: Ponape (MCZ): Koror, Palau Islands
(ANSP). Japan: Oshima Osumi (MCZ); Chishima, Okinawa, Ryukyu Islands (J.
Schwengel). PuitippinE Istanps: Lubang (MCZ). Ausrratia: Debris Bay, Cape
Grafton, near Cairns, Queensland (J. K. Howard): Murray Island, Great Barrier Reef,
Queensland (MCZ). Ixnp1an Ocean Isuanps: Ceylon; Zanzibar (both MCZ). Porrv-
GUESE East Arrica: Bazaruto Island, Bazaruto Bay (J. K. Howard).

Subgenus Gutturnium Moérch

Gutturnium Morch 1852, Catalogus Conchyliorum Comes de Yoldi, p. 109.

Type species, 7'riton tuberosum Lamarck [| = Distorsio muricina Réding), subsequent
designation, Dall 1904.

Shell medium in size and gray to brown in color. Sculpture consisting of a few varices,
with one to four knobbed axial ridges. Spiral sculpture of numerous beaded cords, with
finer cords between them. Wohorls convex. Siphonal canal narrow, moderately extended

B} 9e Y . aie oa toale P . . .
Plate 126. Cymatium muricinum Riding. Figs. 1-2. Caleton de Don Bruno, Cienfuegos, Cuba (2.16x). Fig.

3. Savu Savu Bay, Vanua Levu Island, Fiji Islands (2.16x).

Western Atlantic JOHNSONIA, No. 36 225

and directed upward. Aperture subcircular and having a broad parietal shield. Perios-
tracum thin and deciduous. Operculum unguiculate with an apical nucleus and sculp-
tured with concentric growth lines.

There is only one species in this subgenus in the Western Atlantic.

Cymatium (Gutturnium) muricinum Réding
Plate 112, figs. 5-6; Plate 113, fig. 8; Plate 126, figs. 1-3; Plate 127

Distorsio muricina Réding 1798, Museum Boltenianum, p. 133 [refers to Martini (1) 3, pl. 112, figs. 1050-
1051].

Murex pyrum Gmelin 1791, Systema Naturae, ed. 13, p. 3535 [in part, reference to Martini (1) 3, pl. 112,
figs. 1050-1051, only].

Triton tuberosum Lamarck 1822, Histoire Naturelle des Animaux sans Vertébres 7, p. 185 (l’Océan des
Grandes Indes). [Refers to Martini (1) 3, pl. 112, figs. 1050-1051. |

Triton pyriformis Conrad 1849, Journal Academy Natural Sciences Philadelphia (n.s.) 1, p. 211 [refers to
Martini (1) 3, pl. 112, figs. 1050-1051].

Triton productum Gould 1852, United States Exploring Expedition 12, Mollusca, p. 240 (Fiji Islands).

Triton antillarum d’Orbigny 1842 [in] Sagra, Histoire de l’Ie de Cuba, Mollusques 2, p. 161, pl. 23, fig.
20 (Cuba).

Litiopa obesa C. B. Adams 1850, Contributions to Conchology, no. 5, p. 71 (Jamaica) [embryonic shell].

Description. Shell medium in size, reaching about 75 mm. (about 3 inches) in length,
solid, imperforate and strongly sculptured. Color dull grayish brown, with occasional
banded specimens having broad bands of dark reddish brown; often these bands are visi-
ble only in transmitted light. Lip and parietal shield white to light ivory and highly
glazed. Interior of aperture a dark reddish brown. Post-embryonic whorls 5 to 7, and
strongly convex. Spire elevated and produced at an angle of about 60°. Aperture sub-
circular with the outer lip greatly thickened when a varix is produced. Parietal lip glazed
to form a rather broad shield. There are 4 or 5 inconspicuous plicae at the base of the
columella. The outer lip has 7 strongly developed crenulations. Siphonal canal extended,
though variable in length, and directed upward at an angle of about 55°. Columella
slightly arched inwardly, its base continuing into the parietal margin of the siphonal
canal. Suture irregular and somewhat indented. Sculpture consisting of numerous spiral
and nodulose cords of unequal strength, and numerous fine, thread-like ridges. Axial
sculpture consisting of 7 or 8 nodulose varices. In addition there are 2 to 8 knobbed ridges
between the varices which give the shell a very rugose appearance. Periostracum produced
in numerous rows of thin low blades, light brown in color and deciduous. Operculum un-
guiculate, its nucleus apical and the surface roughened by numerous fine concentric ridges.

length width
75mm. 37.5mm. Mokuole Island, Kaneohe Bay, Oahu, Hawaiian Islands
if 33 Port Antonio, Jamaica
58 28 Mauritius
49 31 Caleton de Don Bruno, 4 mi. SW of Cienfuegos, Cuba
46 Dayal Savusavu, Vanua Levu, Fiji Islands

Types. The type figures are here selected to be Martini, Conchylien-Cabinet (1) 3,
pl. 112, figs. 1050-1051, for muricinum Réding, tuberosum Lamarck and pyriformis Con-
rad, as indicated in the synonymy, were all based on these figures. The type of antillarum
d’Orbigny is in the British Museum (Natural History). The holotype of productum

JOHNSONIA, No. 36 Cymatium

226

Gould is probably in the United States National Museum. The type locality as given
by Martini is the Coromandel Coast, India. The holotype of L.obesa C. B. Adams is in

the Museum of Comparative Zoology, no. 186594.

Remarks. So far as we can detect there is no appreciable difference between the West-
ern Atlantic and the Indo-Pacific specimens of the species. It is quite variable in size at
maturity and specimens also vary as to the number and development of the nodulose
varices and ridges.

Range. WesveERN ATLANTIC: Bermuda and from Jupiter Inlet, Florida and Vera-
cruz, Mexico throughout the West Indies and south to Estado do Parana, Brasil (Mor-
retes, 1949).

Inpo-Paciric: From the Hawaiian Islands south to the Marquesas, west to the Ry-
ukyu Islands, the Philippines south to Australia and the East Indies and west in the

Indian Ocean to Zanzibar.

Specimens examined. WESTERN ATLANTIC. FLoripa: Jupiter Inlet (FSM); Lake
Worth, Boynton (MCZ; FSM: ANSP; T. McGinty); Boca Raton (J. N. Flowers);
off Fort Lauderdale (MCZ): Key Biscayne (J. K. Howard: R. Work); Grassy Key
(ANSP): Boca Chica Key (D. and N. Schmidt); Pelican Shoals, Key West; Dry Tor-
tugas (both FSM). Bermupa: Castle Harbour (MCZ). Banama IsLanps: North Point,

I}. oF J . o <6 - J- : ~ ons =
Plate 127. Cymatium muricinum Réding. Miragoane, Haiti (to show embryonic whorls, 30x).

Western Atlantic JOHNSONIA, No. 36 297

Elbow Cay, Great Abaco (R. Robertson); Settlement Point, Grand Bahama; Adelaide,
New Providence (both MCZ); Lyford Cay, New Providence (G. & M. Kline); South
Bimini; Little San Salvador; Arthurs ‘Town, Cat Island; Clarence Town, Long Island
(all MCZ); Man Island, Eleuthera (G. and M. Kline). Cusa: Varadero, Matanzas
(ANSP; J. Schwengel); Cardenas, Matanzas (MCZ; J. Finlay); Cayo Francés, Cai-
barién, Las Villas (P. J. Bermudez); Cayo Santa Maria, Camagiiey (R. Humes): Banes,
Oriente; Santiago, Oriente (both C.G. Aguayo); Blue Beach, Guantanamo Naval Base,
Oriente; Caleton de Don Bruno, Cienfuegos (both MCZ). Hispanio.a: Miragoane,
Haiti; Montecristi; Puerto Plata and Santa Barbara de Samana, all Santo Domingo
(all MCZ). Jamaica: Port Antonio (MCZ). Purrro Rico: San Juan (D. Thomas):
north of Veja Baja (MCZ); Arecibo; Aguada (both G. Warmke); Bahia de Afasco;
Punta Guanajibo; Mayagiiez; Ponce (all MCZ; UP; ANSP); Puerto Patillas (G.
Warmke). Vircin IsLanps: St. Thomas; St. John (both MCZ); Guana Island, Tor-
tola (M. W. Dewey); St. Croix (FSM; ANSP). Lesser ANTILLES: Guadeloupe
(ANSP); Barbados(MCZ). CartBBEan IsLANDS: Curacao, Dutch West Indies. Merx-
Ico: Veracruz, Veracruz (T.Pulley; MCZ). Panama: Colén (ANSP).

Inpo-Paciric. Hawaiian Istanps: Mokuole Island, Kaneohe Bay, Oahu; Kawai-
hae, Hawaii; Hilo Bay, Hawaii (all MCZ). Line Istanps: Palmyra Island (MCZ).
BakER IsLaNnD (MCZ). Socitery IsLanps: Punaauia and Arue, Tahiti; Teavaro-
Teaharoa, Moorea (all R. Robertson). Marsuautui Istanps: Ebon Island (MCZ). GIL-
BERT IsLANDS: Apiang(MCZ). Marianas Istanps: Tumon Bay, Guam (MCZ); Apra
Harbor, Guam (W.Old). CaroLine IsLaNps: Ponape (MCZ); Peleliu Island and Koror
Island, Palau Islands (both ANSP). Fis1 IsLanps: Valanga Bay, Vanua Levu; Suva,
Viti Levu (both MCZ); Bega Island (J. Schwengel). NEw CaLeponia (MCZ). NEw
GuINEA: Sowek, Soepiori Island, Schouten Islands, Dutch New Guinea (ANSP).
AvstTRALIA: T'wo Isles, Queensland (E. Grigg). JAPAN: Ookamijima, Taira, Miyako,
Ryukyu Islands; Machinato, Okinawa, Ryukyu Islands (both MCZ). Puivipepine
IsLaNps: Guiuan, Sdmar; Calapan, Mindoro; Lubang; off Aborlan, Palawan (all
MCZ). Motucca IsLanps: Tengah Island, off Bouro Island; Bouro Island; Amboina
Island (all MCZ). INp1an Ocean IsLanps: Hikkadua, Ceylon (ANSP); Mauritius;
Zanzibar (both MCZ). Kenya: Kilifi(E.Grigg). Porrucursr East Arrica: Bazaruto
Island, Bazaruto Bay (J. K. Howard); Mozambique (J. Schwengel).

Subgenus Monoplex Perry
Monoplex Perry 1811, Conchology, pl. 3, fig. 3.

Type species, Monoplex australasiae Perry [| = Cymatium parthenopeum von Salis}, here
selected.

Both Herrmannsen 1846 and Gray 1847 gave M. olearium as the type species of Mon-
oplex, but this name was not included among the species described by Perry for this
subgenus.

Shell medium to moderately large in size and generally yellowish to brownish in color.
Sculpture consisting of large spiral cords and occasionally of varices. The shell is usually
rimately umbilicate. Periostracum heavy, produced in fringed axial blades and usually
deciduous. Operculum unguiculate, with an apical nucleus.

298 JOHNSONIA, No. 36 Cymatium

Cymatium (Monoplex) parthenopeum' von Salis
Plate 110, fig. 4; Plate 112, figs. 7-8; Plate 113, figs. 9-10; Plate 128, figs. 1-3

Triton olearius of authors not the Murex olearium Linné 1758 and 1767. We here select Linné’s reference
to Gualtieri 1742, pl. 50, fig. A to be the type figure of M. oleartum Linné which is an earlier name for Ranella

gigantea Lamarck.
Murex costatus Born 1778, Index Rerum Naturalium Musei Caesarei Vindobonensis 1, p. 295 (locality un-

known); Born 1780, Testacea Musei Caesarei Vindobonensis, p. 297 [refers to Seba 1758, Thesauri 3, Dla ow.
fig. 31], non Murex costatus Pennant 1777.

Murex parthenopeus von Salis 1793” Reisen in versch. Prov. Kénigreich Neapel 1, p. 370, pl. 7, fig. 4;
1795, English translation by Aufrere, p. 462, pl. 7, fig. 4 (Bay of Naples).

Triton succinctum Lamarck 1816, Tableau Encyclopédique et Méthodique 3, pl. 416, fig. 2; Liste, p. 5;
Lamarck 1822, Histoire Naturelle des Animaux sans Vertébres 7, p. 181 (mers de la Nouvelle-Hollande).

Triton americanum d’Orbigny 1842 [in] Sagra, Histoire de l’Ile de Cuba, Mollusques 2, p. 163, pl. 23, fig.

22 (Cuba).
Triton brasilianum Gould 1849, Proceedings Boston Society Natural History 3, p. 142 (Rio de Janeiro,
Brasil); Gould 1856, Appendix, p. 504 of the United States Exploring Expedition 12, pl. 17, fig. 296.
Monoplex australasiae Perry 1811, Conchology, London, pl. 3, fig. 8 (New Holland and Lord Howe’s Island).
Triton (Simpulum) acelivis Hutton 1873, Catalogue of the Marine Mollusca of New Zealand, Wellington,

p. 13, fig. 8 (New Zealand).

Description. Shell large in size, reaching 145 mm. (about 5% inches) in length, rather
heavy in structure, imperforate or with a small umbilical chink and with a pronounced
spiral sculpture. Color brownish yellow, usually fairly uniform, but occasionally with
spiral bands of slightly darker brown which become much darker on the varices. Embry-
onic whorls three, smooth, light straw-yellow in color, extended and slightly tilted from
the remaining whorls. Post-embryonic whorls 7 to 8, convex and slightly shouldered.
Spire moderately extended and produced at an angle of about 55°. Aperture subellipti-
cal, the outer lip margined within by coarse teeth which are grouped in pairs, occasion-
ally having smaller secondary teeth. These palatal teeth are opposite the grooves which are
between the spiral cords. Parietal wall with numerous, irregular, white plicae, the area
between the plicae being a dark reddish brown. There is a single large ridge on the upper
portion of the parietal wall which margins the anal canal. Siphonal canal somewhat vari-
able and curved upward and slightly to the left when viewed dorsally. Columella thick-
ened and continuing as the parietal margin of the siphonal canal. Suture slightly indented.
Sculpture consisting of 5 or 6 broad, low and often nodulose, spiral cords, with numerous
finer cords on the base and on the siphonal canal. In addition, there are numerous fine
spiral threads which appear in the interspaces as well as on the cords themselves. Axial
sculpture consisting of numerous fine growth lines. Varices rather low and usually with
only two, including the lip, showing in fully grown adult specimens. Periostracum rather
thin, a dark golden brown in color and produced in numerous axial blades with a hair-
like fringe, and usually deciduous. Operculum unguiculate with a terminal nucleus and
sculptured with numerous concentric growth lines.

'Parthenope was an old name for the town of Neapolis [| Naples].

* We have seen only the English translation by Anthony Aufrere 1795, ““Travels through various Provinces
of the Kingdom of Naples in 1789,’’ London, 527 pages, 9 plates.

Western Atlantic JOHNSONIA, No. 36 Q¢

29
length width
145 mm, 79mm. Bermuda
109 64 Nagasaki, Japan
96 56 Rio de Janeiro, Brasil
89 SYN. Moreton Bay, Queensland, Australia

Types. The type of Murex costatus Born is in the Zoological Museum, Vienna, Aus-
tria. The type of T7iton americanum d’Orbigny is in the British Museum (Natural His-
tory). A paratype of 7° brasilianum Gould is in the Museum of Comparative Zoology,
no. 191179. We do not know the location of the type specimens of the other names
listed in the synonymy. The type locality is Naples, Italy.

Remarks. The distribution of this species is quite remarkable, existing as it does in
most tropical and warm temperate seas. On the basis of shell morphology we can detect
no consistent differences in the specimens from any of the widely separated localities.
This opinion was held by Watson (1886, p. 391) who reviewed most of the material con-

Plate 128. Cymatium parthenopeum von Salis. Fig. 1. Moreton Bay, Queensland, Australia (natural size).
Fig. 2. Sanibel Island, Florida (natural size). Fig. 3. Rio de Janeiro, Brasil (to show embryonic whorls, 8x).

230 JOHNSONIA, No. 36 Cymatium

tained in several of the European museums. There may, of course, be consistent differ-
ences in the soft anatomy, but this study still remains to be done.

Our records are too few to determine whether or not there is a continuous population
between the end localities in any one region. For example, Nicklés (1950, p. 86) states
it occurs from Morocco and south to the Cape of Good Hope, but he neither lists any
localities nor indicates upon how much material this statement was based. We possess
material from Japan, Australia and New Zealand, but have no material from between
these very widely separated localities. It is not known to occur in the Eastern Pacific.

The well-known name of M. costatus Born must give way to C. parthenopeum von
Salis because of an earlier use of Murex costatus by Pennant in 1777 as indicated in the
synonymy above. However, the name of von Salis has been used generally by Europeans
for this species.

This species apparently is rarely found alive in the intertidal area, but it does occur
from just below low water to depths of at least 35 fathoms.

De Gregorio (1884, Bullettino della Societa Malacologica Italiana 10, p. 96) described
two living and one fossil varieties of parthenopeum trom Sicily.

Range. EKastERN ATLANTIC: Western Mediterranean, the Azores and south to the
Union of South Africa.

WestrerRN ATLANTIC: Bermuda, Florida and Mexico, the West Indies and south to
Brasil.

Invo-Paciric: Japan, eastern Australia, northern New Zealand and Portuguese
East Africa.

Specimens examined. FLORIDA: south end of Lake Worth; Jupiter Inlet, Palm Beach
Co. (both FSM); Bear Cut, Key Biscayne (R. Work); Dry Tortugas (R. Merrill; FSM);
Naples (MCZ); Sanibel Island (J. Schwengel); St. Andrews Bay, Panama City (R.
Work). Texas: off Port Isabel in 25 fathoms (H. and K. Johnstone). Mexico: off
Laguna Madre, Tamaulipas in 82 to 85 fathoms (H. Hildebrand); 15 miles N of Tecolutla,
Veracruz (T. Pulley); Campeche (H. Hildebrand). BErmupa: (MCZ). Cuspa: Matan-
zas Bay and Varadero Beach, Matanzas (both J. Finlay). Purrro Rico: Mona Island;
Punta Algarrobo (both G.Warmke). VirGin Istanps: St. Thomas (ANSP); Tortola
(MCZ; ANSP). Lessrr AnrTILLES: Barbados; Trinidad (both MCZ). Brasit: Praia
de Leste, Ilha Guaiba, Est. de Rio de Janeiro (de Oliveira); Rio de Janeiro (MCZ);
Praia de Urea, Districto Federal; Manguinhos, Ilha de Itaparica, Est. de Bahia (both
de Oliveira).

EASTERN ATLANTIC. FRANCE: Marseilles (MCZ). [raty: Catania, Sicily (W. Old).
ALGERIA: Gulf of Oran in 19 fathoms (MCZ). Azores: Fayal (MCZ). SENEGAL: Dakar
(MCZ). Sourn Arrica: Jeffreys Bay (D.H. Kennelly); Port Alfred (J.Schwengel).

Inpo-Paciric. NEw ZEALAND: Lauranga Harbour; Hen and Chicken Reef (both
MCZ); Pana, Parengarenga Harbour (A. W. B. Powell). Ausrratia: Moreton Bay,
Queensland (MCZ; J.Schwengel); Narooma, New South Wales (MCZ): Sydney, New
South Wales (J.Schwengel); Ulladulla, New South Wales (E.Grigg). Japan: Awaji;
Kishiu; Misaki, Sagami;. Yenosima; Tokyo Bay; Koka Shima (all MCZ); Nagasaki
(ANSP). Porrucurst East Arrica: Bazaruto Island, Bazaruto Bay (J. K. Howard).

Western Atlantic JOHNSONIA, No. 36 231

Subgenus Cymatium Réding
Cymatium Réding 1798, Museum Boltenianum, p. 129.

Type species, Murex femorale Linné, subsequent designation, Dall 1904.

Shell large, triangular in cross section, and generally a rich golden brown in color.
Sculpture consisting of large knobs and spiral cords, in addition to having three or more
well-developed varices. Both palatal denticles and parietal plicae usually well-developed.
Generally with a depressed area at the beginning of the siphonal canal. Siphonal canal
usually extended and curved upwardly. Periostracum thin and slightly roughened over
most of the surface, but producing high, thin, axial blades which are fringed. Opercu-
lum small for the size of the aperture, unguiculate, with a terminal nucleus and sculp-
tured with rather coarse concentric ridges.

Daa an » ~
Plate 129. Cymatium femorale Linné. Fig. 1. Great Abaco, Bahama Islands (slightly reduced). Fig. 2. Bear
Cut, Miami, Florida (slightly enlarged). Fig. 3. French Beach, Varadero, Matanzas, Cuba (to show em-

bryonic whorls, 12.4x).

232 JOHNSONIA, No. 36 Cymatium

Only one member of this subgenus is found in the Western Atlantic, with a single
species in the Eastern Pacific; the remaining few species are from the Indo-Pacific area.

Cymatium (Cymatium) femorale Linné
Plate 110, fig. 1; Plate 112, figs. 9-10; Plate 113, fig. 11; Plate 129, figs. 1-3
Murex femorale Linné 1758, Systema Naturae, ed. 10, p. 749 (O. Asiatico); Hanley 1855, Ipsa Linnaei
Conchylia, p. 287.
Lotorium lotor Montfort 1810, Conchyliologie Systématique 2, p. 583 (cdtes d’ Afrique).
Septa triangularis Perry 1811, Conchology, pl. 14, fig. 6 (Southern Ocean).
Triton lotorium Lamarck 1816, Tableau Encyclopédique et Méthodique, Liste, p. 5, Atlas 3, pl. 415, fig. 2.

Description. Shell large, reaching 220 mm. (about 84 inches) in length, strong but not
heavy, imperforate and sculptured. Color a golden brown to light reddish brown with
alternating bands of brown and white on the varices. Whorls 8 or 9, convex, somewhat
flattened dorso-ventrally and slightly shouldered. Spire somewhat extended. Aperture
auricular in shape, the parietal wall consisting of a thin glaze; within there may be 1 to
4 or 5 small relatively inconspicuous plicae. Certain of these appear to be independently
produced while others are nothing more than the glaze over the spiral cords. Outer lip
rolled inwardly to form a varix. Columella irregular; it may be nearly straight or strongly
arched inwardly and to the left. Siphonal canal somewhat extended and curved upwardly.
Suture relatively indistinct. Sculpture consisting usually of five varices (lip included),
visible in the adult. The varices are knobbed, the knobs being the high points where the
spiral cords pass over them. There are 6 to 8 rather heavy, conspicuous spiral cords, the
largest and heaviest at the shoulder. In addition, this shoulder cord may be irregularly
nodulose and occasionally smaller nodules are present on the cords below the shoulder
cord. Between the cords there are numerous spiral threads and these are crossed by
somewhat finer axial threads. Operculum unguiculate and relatively small for the size of
the aperture. It has a marginal nucleus and the growth ridges are concentrically devel-
oped. Periostracum thin, light but rough, somewhat foliated, yellowish brown in color
and deciduous. Occasional specimens are found with several axial, fringed blades of peri-
ostracum which correspond with the axial ribs of the shell.

length width
212mm. 115mm. Gulf of Mexico
176 74 Savannah Sound, Eleuthera Island, Bahama Islands
153 105 Cayo Francés, Caibarién, Las Villas, Cuba

Types. The holotype of Murex femorale Linné, according to Hanley (1855) is in the
Linnean Collection of the Linnean Society, London. We here restrict the type locality
to Jamaica, the locality given by Lister in his Historiae Methodicae Conchylorum on
plate 941 to which Linné referred.

Remarks. ‘This is a species of rather wide distribution throughout the West Indian
region. Specimens are not rare, yet they never appear to be common at any one locality.
They are usually found a little beyond low water in areas with a sandy bottom where
sea grass is abundant. Dall (1889, p. 182) has given Cedar Keys as the northern limit of
this species on the west coast of Florida. This appears to be an error, however, as the
species probably does not occur, at least alive, north of Key West on the west coast of
Florida.

Western Atlantic JOHNSONIA, No. 36 233

Range. Bermuda, southeastern Florida and the lower Florida Keys, throughout the
West Indies, and from central Mexico south to Bahia, Brasil.

Specimens examined. FLortpaA: Melbourne, Brevard Co. (FSM); Bear Cut, Key Bis-
cayne, Miami (R. Work; H. and K. Johnstone); Ragged Keys, Biscayne Bay (MCZ);
Key West (ANSP; FSM); Tortugas (J. Miller). BeErmupa: Castle Harbour (dredged
dead, H.B. Moore). Banama Istanps: High Rock, Grand Bahama(MCZ); Elbow Cay,
Great Abaco (G. Kline; R. Robertson); Nixon’s Harbour, South Bimini, Bimini Islands:
Hog Island, New Providence (both MCZ); Dick’s Point, Nassau, New Providence
(G. Kline); Spanish Wells, Eleuthera (MCZ; J. Schwengel); The Current, Eleuthera
(G. and M. Kline); Sandy Point, Savannah Sound, Eleuthera; Little San Salvador:
Orange Creek, Cat Island; Arthurs Town, Cat Island; Wemyss, Long Island; Matthew
Town, Great Inagua (all MCZ). Cuna: near Habana (R.Humes): Cayo Francés, Cai-
barién, Las Villas (Museo Poey); Varadero, Cardenas, Matanzas (ANSP; J. Finlay);
Fishpoint, Guantanamo Bay (MCZ). Hispaniota: Cap Haitien, Haiti (W. J. Eyer-
dam); Montecristi, Santo Domingo; Puerto Plata, Santo Domingo (both MCZ).
JAMAICA: Whitehouse (J. K. Howard). PurErro Rico: Catafio; Arecibo (both G.
Warmke); Punta Borinquén (A. Phares); Rinecén; Bahia de Afiasco; Mayagiiez; En-
senada; Puerto Patillas (all G. Warmke); Punta Guanajibo; Punta Algarrobo (both
UPR). Viren Istanps: Guana Island, Tortola (M. Dewey); St. Thomas (MCZ;
ANSP); St. Johns (MCZ); St. Croix (FSM). Lrssrr ANTILLES: Barbados (MCZ;
ANSP); English Harbour, Antigua (K. Anderson). CarrIBBEAN IsLANDS: Oranjestad
Harbour, Aruba (D. P. Barnes). Mexico: Tampico, Tamaulipas; 15 miles N of Teco-
lutla, Veracruz (both T. Pulley). Nicaracua: Bluefields (MCZ). Panama: Panama
(Atlantic) (MCZ). Brasi.: Itapagipe, Salvador, Bahia (H. L. Lopes).

Cymatium (Cymatium) tigrinum Broderip
Plate 130, figs. 1-2

Triton tigrinus Broderip 1833, Proceedings Zoological Society London, p.5(in America Centrali (Guacomayo) ).

Description. Shell large, reaching 185 mm. (about 7$ inches) in length, strong and

fairly heavy, imperforate and sculptured. Color a light reddish brown to yellow, overlaid
by arather dark reddish brown, roughened periostracum. Varices dark brownish red be-
tween the spiral cords. This color is also present inside the outer lip. Base of the colu-
mella may also be stained a dark mahogany-brown. Whorls 8 to 84, convex and flat-
tened dorso-ventrally. Spire extended. Aperture subelliptical. Outer lip thickened when
the heavy varix is present. Inner lip consisting of a somewhat heavy glaze on the parietal
area. Outer lip rolled inwardly to form a varix. Columella straight to slightly curved.
Siphonal canal somewhat extended and curved upwardly. Sculpture consisting usually
of five varices (lip included) which are visible in the adult. These varices are doubled in
all specimens which we have seen, except for the last varix. The varices are knobbed,
the knobs being the high points where the spiral cords pass over the varices. ‘There are
8 or 9 conspicuous spiral cords; the largest may possess 1 or 2 rather large nodules. In
between the cords there are numerous and relatively indistinct spiral threads. Apical
sculpture consisting mainly of very fine growth lines. Periostracum rather heavy, rough-
ened, particularly at the varices and deciduous. Operculum unknown.

234 JOHNSONIA, No. 36 Cymatium

length width
185 mm. 101mm. Acapuleo, Mexico
164 86 Corinto, Nicaragua
111 65 Panama

Types. The holotype of 7. tigrinus Broderip is probably in the British Museum
(Natural History). The type locality is Guacomayo, Central America. We have been
unable to locate any coastal village by this name.

Remarks. This is a very distinct species and apparently not closely related to any other
known species. We have added it here only as representing the Eastern Pacific analogue
of the Western Atlantic C. femorale. The two species, however, are not closely related.

This appears to be a very rare species to Judge by the few specimens that are to be
found in our museums. The range we give below is probably not the limitations of this
species. It may very well extend south at least to northern Ecuador.

Plate 130. Cymatium tigrinum Broderip. Figs. 1-2. Costa Rica. (California Academy of Sciences, no. 33079,

natural size.) Photograph through the kindness of Dr. L. G. Hertlein.

Western Atlantic JOHNSONIA, No. 36 235

Range. From La Paz, Baja California (Pilsbry and Lowe 1932) south to Panama.

Specimens examined. Mexico: Acapulco (MCZ; San Diego Soc. Nat. Hist.): Est.
Oaxaca (Stanford Univ.). Nicaracua: Corinto (San Diego Soc. Nat. Hist.). Cosra
Rica: (California Acad. Science; USNM). Panama: (MCZ; ANSP; USNM). Ve-
nado Island (J. Schwengel; Mrs. J. D. Elliott); Balboa, Canal Zone (W. Old).

Genus Distorsio Réding

Distorsio Riding 1798, Museum Boltenianum, p. 133.

Distortrix Link 1807, Beschreibung der Naturalien-Sammlung der Universitat zu Rostock, p. 122 (type spe-
cies, Distortrix anus Linné | = Murex anus Linné]}, subsequent designation, Dall 1904).

Persona Denys de Montfort 1810, Conchyliologie Systématique 2, p. 603 (type species, Persona anus Linné
[ = Murer anus Linné]|, monotypic).

Distorta Perry 1811, Conchology, pl. 10, fig. 2 (type species, Distorta rotunda Perry |= Murex anus Linné],
subsequent designation, Puffer and Emerson 1953).

Distortio ‘Bolten’ Gray 1847, Proceedings Zoological Society London, p. 133 [correction for Distorsio Réding].

Distortic ‘Link’ Paetel 1875, Familien- und Gattungsnamen der Mollusken, Berlin, p. 71 [error for Dis-
tortrix Link].

Persoma ‘Montfort’ Paetel 1875, Familien- und Gattungsnamen der Mollusken, Berlin, p. 157 [error for
Persona Montfort }.

Distorsus ‘Bolten’ Paetel 1888, Catalog der Conchylien-Sammlung 1, p. 103 [error for Distorsio Réding}.

Distorsia ‘Riding’ Pilsbry 1922, Proceedings Academy Natural Sciences 73, p. 359 [error for Distorsio

Réding].
Type species, Murex anus Linné, subsequent designation, J. E. Gray 1847.

Winckworth in 1945 apparently overlooked Gray’s type designation for this genus in
1847. Even though Gray corrected the spelling to Distortio there is no question that he
was referring to Distorsio ‘Bolten’ Réding. Pilsbry (1922) has been credited with the
type designation.

Shell ranging from 25 mm. (1 inch) to 75 mm. (about 3 inches) in length, imperforate,
subglobose to elongate, and rather solid in structure. Color ranging from a grayish white
to mottled brown. Whorls very irregular and distorted. Sculpture consisting -of spiral
cords and axial costae. Parietal shields thin but extended and portions of previous shields
remain on earlier whorls producing the varices. Aperture auricular in appearance being
complicated by numerous palatal denticles and parietal plicae. ‘The denticles and plicae
produced at earlier stages of growth are not absorbed as the shell grows but remain as
internal structures in the shell as shown in Plate 131. Because of this apertural armature
the shell is twisted and bulges to compensate for the space occupied by these internal
denticulations.

Species in this genus are found in most tropical portions of the world and range from
low water to depths of about 300 fathoms. According to Woodring (1928, p. 299) the
earliest Distorsio s.s. appeared in the Byram marl, Upper Oligocene of Mississippi.

Some of our recent species are exceedingly close, if not identical with certain fossil
forms. This is particularly true of D. megintyi Emerson and Puffer, and D. decussatus
simillimus Sowerby as figured by Woodring (1928, plate 18, fig. 9 and plate 19, fig. 1).
The other specimens figured by Woodring (plate 18, figs. 7-8) appear to be clathrata
Lamarck.

236 JOHNSONIA, No. 36 Distorsio

The opercula of all species so far examined are quite irregular, small for the size of the
shell and often broken, undoubtedly a result of the great irregularity of the aperture.
The nucleus of the operculum is submarginal. The general proportions and shape of the
operculum and the muscle scars appear to be quite variable as shown on plate 1382.

The radula is taenioglossate with a relatively small central tooth, a broad and denti-
culate lateral tooth and two subequal marginal teeth.

Subgenus Distorsio Réding
Distorsio Réding 1798, Museum Boltenianum, p. 133.

Type species, M. anus Linné, subsequent designation Gray 1847.

This subgenus contains but a single species and is characterized by the shell producing
an exceedingly large parietal shield which extends well above the upper limits of the
outer lip and covers the two preceding whorls. The early shields are not absorbed but
remain as thin blade-like varices. Siphonal canal is nearly vertical and not visible from
the apertural side.

Distorsio anus Linné, the only species now known to belong in the subgenus Distorsio,
is wide-ranging throughout most of the tropical Indo-Pacific.

Subgenus Rhysema,’ new subgenus

Shell ranging in size from 25 mm. (1 inch) to 90 mm. (about 83 inches). Color rang-
ing from grayish white or light yellow to pinkish. Parietal shield broad but not extend-
ing above the upper limit of the outer lip. Sculpture consisting of spiral cords and axial
costae, both being about equal in coarseness. Siphonal canal curved upwards but not
vertical.

This subgenus differs from typical Distorsio by lacking the extended parietal shield
and by not having the siphonal canal vertical.

Species in this subgenus occur in nearly all tropical and subtropical seas.

Type species, Triton clathratus Lamarck.

Distorsio (Rhysema) clathrata Lamarck
Plates 131; 132, figs. 2-8; 1383

Triton clathratum Lamarck 1816, Tableau Encyclopédique et Méthodique, Liste, p. 4, Atlas 3, pl. 413,
fig. 4a-b; Lamarck 1822, Animaux sans Vertébres 7, p. 186 [reference to the Tableau only] (mers de
l’Amérique Méridionale).

Distortrix reticulata ‘Link’ Dall 1889, Bulletin Museum Comparative Zoology 18, p. 221 [in part]; non
D. reticulata Link 1807 | =D. reticulata Réding 1798].

Description. Shell medium in size, reaching 77 mm. (about 3 inches) in length, rather
solid, imperforate and strongly sculptured. Color white with a diffusion of yellowish or
pinkish brown. Whorls 10, convex and irregular. Spire extended and produced at an
angle of about 45°. Aperture auricular in shape, the outer lip being somewhat thickened
and with 10 denticles, the third below the anal canal being the largest, while the smallest

‘From the Greek, wrinkle; pucker.

Western Atlantic JOHNSONIA, No. 36 237

one margins the siphonal canal. The largest denticle is opposite the deep parietal embay-
ment. Inner lip consisting of numerous plicae. There are two large parietal plicae which
margin the anal canal. On the lower portion of the columellar area there are 13 or 14 pli-
cae. These are smallest near the siphonal canal, becoming larger posteriorly and project
outward to form the anterior margin of the parietal embayment. In the parietal embay-
ment the spiral cords are slightly thickened. A thin parietal shield is developed which is
margined on the parietal side by glazing over the back of the thin varix and continuing
as aridge both above on the upper portion of the whorl and below to the mid area of the
siphonal canal. Columella nearly straight but complicated by the numerous plicae. These
plicae follow back to the early whorls. The denticles which were produced on early lips
are not absorbed but remain within the shell as shown in Plate 131. Siphonal canal mod-
erately short and narrow, deflected slightly toward the outer lip and turned slightly up-
ward. Anal canal formed by the uppermost palatal denticle and the two parietal plicae.
Suture slightly impressed, irregular and occasionally obscure. Spiral sculpture consisting
of numerous cords interspaced with numerous and very fine spiral threads. Axial sculp-
ture consisting of numerous ridges which cross the spiral cords to form a reticulated pat-
tern. Where the spiral cords and axial ridges cross they produce small knobs. There are
7 to 9 varices. Operculum small for the size of the shell due to the distorted aperture.
It is unguiculate in shape, has a submarginal nucleus and is sculptured with numerous
concentric growth lines. Periostracum thin, yellowish brown in color, finely reticulate,
and having numerous, fine hair-like processes over the entire surface and coarse hair-like
processes on the knobs. The embryonic shell has 3 whorls which are smooth, glass-like
in appearance and light amber in color.

Plate 131. Distorsio clathrata Lamarck, Tuxpan, Veracruz, Mexico (1.5x). A sectioned shell to show reten-
tion of the early varices and apertural armature.

238 JOHNSONIA, No. 36 Distorsio

length width
77 mm. 43mm. off Key West, Florida
70 38 Bahia de Campeche, Mexico
67 34 West of Tortugas, Florida
62 34 Puerto Plata, Santo Domingo

Types. The present location of the holotype of Dzestorsio clathrata Lamarck is un-
known to us. It is probably in the Paris Museum. The type locality as given by Lamarck

Plate 132. Radulae and opercula of Distorsio. Fig. 1. Radula of Distorsio anus Linné, Amboyna, Molucca
Islands. Fig. 2. Radula of Distorsio clathrata Lamarck, Punta Algarrobo, Puerto Rico. Figs. 3-4. Opercu-
lum of Distorsio clathrata Lamarck from the same locality. Figs. 5-6. Distorsio clathrata Lamarck, 20 miles
NE of North Pass, Mississippi Delta in 100 fathoms. Figs. 7-8. Distorsio clathrata Lamarck, SW of Des-
tin, Florida in 18 to 20 fathoms. Figs. 9-10. Distorsio megintyi Emersonand Puffer, SSW of Sombrero Key,
Key Vaca, Florida. Figs. 11-12. Distorsio anus Linné, Pwakuu Island, West Zanzibar in 11 to 18 fathoms,
a young and broken operculum. Figs. 13-14. Distorsio anus Linné, Amboyna, Molucca Islands.

—— 2

Western Atlantic JOHNSONIA, No. 36 239

is ‘‘mers de 1’ Amérique Méridionale.’’ We here restrict the type locality to the Bay of
Campeche, Mexico, an area from which Lamarck had received much material.

Remarks. The two species in this genus found in the Western Atlantic are quite easily
differentiated. The adult specimens of D. clathrata are nearly twice as large as those of
D. megintyi. In addition, D. clathrata has two parietal plicae near the siphonal canal
while there is only one in meginty. There are 13 or 14 columellar plicae which extend well
down the siphonal canal in D. clathrata but only 9 in megintyi and these stop abruptly
about half way down the siphonal canal. In fully adult specimens of D. megintyi one of
the spiral cords becomes thickened and forms a plica in the center of the parietal embay-
ment. There is only a slight thickening of the spiral cords in this area in D. clathrata.
The irregular coiling of the whorls is far more accentuated in D. megintyi than in clath-
rata and the siphonal canal is much shorter.

Dall in 1889 (loc. cit.) was the first to note that there were two different forms of Dis-
torsio in the Western Atlantic. However, these were not named as new but considered
varieties of his reticulata, a name now applied to an Indo-Pacific species. Olsson and
McGinty first described the deeper water form as Distorsio constricta floridana but un-
fortunately the name floridana was preoccupied and so it was renamed megintyi by
Emerson and Puffer.

Distorsio clathrata Lamarck is generally found from just below low water to depths
of about 30 fathoms though we have examined two lots supposedly taken from depths
of about 100 fathoms.

Range. From Cape Hatteras, North Carolina (Dall 1889, Bulletin United States
National Museum, no. 87, p. 182) where it occurs occasionally and from Lake Worth,
Florida, the northern Gulf of Mexico south to British Guiana. It is found throughout
the West Indies.

Specimens examined. Fitoripa: South Inlet, Lake Worth (MCZ; ANSP); Lake
Worth, Boynton (T.McGinty); off Palm Beach in 20 fathoms (ANSP); off Sombrero
Key, Key Vaca in 30 to 60 fathoms (MCZ; ANSP); off Key West in 17 fathoms (H.
and K. Johnstone); W of Dry Tortugas in 10 to 12 fathoms (J.S.Schwengel); 30 miles
NE of Dry Tortugas (ANSP); 18 miles SW of Destin in 18 to 20 fathoms (T. McGinty);
off Fort Walton in 20 fathoms (MCZ). ALAaBamMa: 20 miles off Petit Bois Island in 10
fathoms; 45 miles off Bayou La Batre (both H. and K. Johnstone); Mississippi: 2 to 38
miles off Horn Island in 12 to 14 fathoms (H. and K. Johnstone). Louis1ana: 20 miles
NE of North Pass, Mississippi Delta in 100 fathoms; off Pass Saluda Light (both H.
and K. Johnstone). Texas: off Aransas Pass (ANSP); Mustang Island (J. W. Hedg-
peth); off Port Isabel (L. A. Weisenhaus); SE of Pass Cavallo and E of St. Joseph
Island (both H. Hildebrand); Brownsville (ANSP). Banama Isutanps: Bullocks Har-
bour, Great Harbour Cay, Berry Islands (G. and M. Kline). Cuna: Matanzas Bay,
Matanzas (C. J. Finlay). Hispanrota: Puerto Plata, Santo Domingo (MCZ). Purrro
Rico: Aguadilla (A. Phares); Bahia de Afiasco (MCZ); Punta Algarrobo; Mayagiiez
(both G. L. Warmke). VirGin IsLanps: St. Thomas(ANSP). Mexico: Bahia de Cam-
peche (MCZ; Museo Poey; H. and K. Johnstone); Veracruz, Veracruz (ANSP; M.E.
Bourgeois); Tuxpan, Veracruz (M. E. Bourgeois); Tampico, Tamaulipas (‘T. Pulley);

240 JOHNSONIA, No. 36 Distorsio

Isla Mujeres, Yucatan (C. G. Aguayo): Punta Frontera, Tabasco (ANSP). Panama:
2 miles off Colén (W. J. Clarke). Cotompra: Cartagena (MCZ). VENEzvELA: Guanta
(ANSP). Brirish Guiana: 4 miles E of Georgetown (H. G. Kugler).

ul Sen oO the ia =
Plate 1338. Distorsio clathrata Lamarck. Figs. 1 and 3. West of Dry Tortugas, Florida in 10 to 20 fathoms
(1.3x). Fig. 2. Boynton, Lake Worth, Florida (1.3x). Fig. 4. Off Fort Walton, Florida in 13 to 19 fathoms,
showing embryonic whorls (18x).

Distorsio (Rhysema) megintyi Emerson and Puffer
Plates 132, figs. 9-10; 134

Distortrix reticulata var. clathrata ‘Lamarck’ Dall 1889, Bulletin Museum Comparative Zoology 18, pp. 221-
229 (Lesser Antilles): non clathrata Lamarck 1816.

Distorsio constricta floridana Olsson and McGinty 1951, Nautilus 65, p. 27, pl. 1, figs. 5, 6, 9 (off Palm
Beach, Florida, in 30-40 fathoms); non D. floridana Gardner 1947 (Miocene).

Distorsio megintyi Emerson and Puffer 1953, Proceedings Biological Society of Washington 66, p. 101 [new
name for D. constricta floridana Olsson and McGinty 1951; non Gardner 1947 |.

Description. Shell small to medium in size, reaching 46 mm. (about 1# inches) in
length, solid, imperforate and strongly sculptured. Color white to pale yellowish brown.
Whorls 9 to 10, convex and irregular. Spire moderately extended and produced at an
angle of about 45°. Aperture auricular in shape, the outer lip somewhat thickened and
with 8 or 9 denticles, the third below the anal canal being the largest, while the smallest
one margins the anal canal. The largest denticle is opposite the deep parietal embay-
ment. Inner lip consisting of numerous plicae. There is a single large plica margining
the anal canal. On the lower portion of the columella area there are 9 plicae which are
smallest near the siphonal canal, becoming larger posteriorly and projecting outward to
form the anterior margin of the parietal embayment. In the center of the parietal em-
bayment there is a single large plica which is developed upon one of the spiral ridges. A

Western Atlantic JOHNSONIA, No. 36 241

thin parietal shield is developed which is free at its parietal margin and curved away from
the shell. Columella nearly straight but complicated by the numerous plicae which con-
tinue back to the early whorls. Siphonal canal short, curved upwardly and toward the
outer lip. Anal canal formed by the palatal denticle and the single parietal plica. Suture
slightly impressed, irregular and occasionally obscure. Spiral sculpture consisting of
numerous cords interspaced with numerous and very fine spiral threads. Axial sculpture
consisting of numerous ridges which cross the spiral cords forming a reticulated pattern.
In many specimens the axial ridges are more prominent. Where the spiral cords and
axial ridges cross each other, small knobs are produced. There are 5 or 6 varices. Peri-
ostracum thin, yellowish brown in color, finely reticulate, having numerous very fine
hair-like processes over the entire surface with a single large periostracal hair on each
knob. Embryonic shell having 8 whorls which are smooth, glass-like in appearance and
light amber in color. Operculum small for the size of the shell, unguiculate in shape,
probably with a submarginal nucleus, and sculptured with concentric growth lines.

Plate 134. Distorsio megintyi Emerson and Puffer. Fig. 1. SW of Sombrero Light, Sombrero Key, Florida
(1.3x). Fig. 2. Off Barbados, Lesser Antilles, Hassler voyage in 100 fathoms (1.38x). Fig. 3. Off Barba-
dos, Lesser Antilles, Blake voyage in 82 fathoms (1.3x). Fig. 4. Off Barbados, Lesser Antilles, Blake voy-
age in 100 fathoms, showing embryonic whorls (16x). Fig. 5. Holotype of Distorsio floridana Olsson and
McGinty (=D. megintyi Emerson and Puffer), off Palm Beach, Florida in 30 to 40 fathoms (4.4x).

242 JOHNSONIA, No. 36 Distorsio
length width
46 mm. 27mm. Off Barbados in 100 fathoms (Hassler Voyage)
43 26 Off Barbados in 82 fathoms (Blake Voyage)
32.5 20 Off Key Vaca, Florida in 70 fathoms
27.8 lao Holotype of D. constricta floridana Olsson and McGinty

Types. The holotype of D. constricta foridana Olsson and McGinty [= D. mcgintyi
Emerson and Puffer] is in the Academy of Natural Sciences Philadelphia, no. 187684.
The type locality is ‘Off Palm Beach, Florida in 30 to 40 fathoms. ”’

Remarks. See Remarks under D. clathrata Lamarck. This is a deep water species, hav-
ing been taken in depths of 30 to about 200 fathoms. It is most closely related to D.
constricta Broderip of the Eastern Pacific from which it differs by having a pronounced
plica in the parietal embayment and only a single parietal plica at the anal canal. In
constricta the plica in the parietal embayment is small or wanting and there may be two
or more plicae on the parietal wall at the anal canal. Both species are very irregularly
coiled and otherwise quite similar.

Range. From off Palm Beach, Florida, the Florida Keys and south through the
West Indies to the Island of Barbados.

Specimens examined, Fitoripa: off Palm Beach in 30, 40, 58 and 70 fathoms (T.
McGinty; MCZ; ANSP); 34 miles NE of Pacific Reef off Elhott Key in 66 fathoms
(MCZ); off Carysfort Light, Key Largo (MCZ); 55 miles SE of The Elbow, Key Largo
in 66 fathoms (MCZ); off Sombrero Light, Key Vaca in 40 fathoms (T. McGinty;
MCZ: ANSP); off Key Vaca in 70 fathoms (T. McGinty); off Pensacola in 45 fathoms
(ANSP). Cusa: Bahia de Matanzas, Matanzas in 120 fathoms (C. J. Finlay). Lesser
ANTILLES: off Guadeloupe, Blake Station 164 in 150 fathoms; off Dominica, Blake
Station 177, in 118 fathoms; off Barbados, Blake Station 278, in 103 fathoms: off Bar-
bados, Hassler Stations in 82 and 100 fathoms; off Grenada, Blake Station 247 in 170
fathoms; off Grenada, Blake Station 262 in 92 fathoms (all MCZ).

Notes

Under Cymatium caribbaeum Clench and Turner (p. 206) we discussed the confusion of
names that existed for cynocephalum Lamarck and cingulatum Lamarck. The following
is at least a partial list of the synonyms of cynocephalum in the Eastern Pacific. This spe-
cies appears to be closely related to C. poulsenitti Mérch of the Western Atlantic.

Cymatium cynocephalum Lamarck
Plate 135, figs. 1-2

Triton cynocephalum Lamarck 1816, Tableau Encyclopédique et Méthodique, Liste, p. 5; Atlas, 3, pl. 422,
fig. 3; Lamarck 1822, Histoire Naturelle des Animaux sans Vertébres 7, p. 184 (locality unknown); non
Triton cynocephalum ‘Lamarck’ Kiener 1842, and of subsequent authors.

Cassidaria cingulata Lamarck 1822, Animaux sans Vertébres 7, p. 216 (no locality given).

Fusus wiegmanni Anton 1839, Verzeichniss der Conchylien, p. 77 (no locality given).

Western Atlantic JOHNSONIA, No. 36 243

Triton undosum Kiener 1842, Iconographie des Coquilles Vivantes 7, Triton, p. 44, pl. 6, fig. 2 (locality
unknown) [substitute name for Cassidaria cingulata Lamarck 1822).

Triton chemnitzii “Gray’ Reeve 1844, Conchologia Iconica 2, Triton, pl. 11, fig. 37 (Panama); non Triton
chemnitzi Gray 1839.

Cassidaria setosa ‘Hinds’ Reeve 1844, Conchologia Iconica, 2, Triton, pl. 11, fig. 37 [nomen nudum, in the

synonymy of Triton chemnitzii Reeve].

TOS eas

Plate 135. Cymatium cynocephalum Lamarck. Fig. 1. Bahia Magdalena, Baja California, Mexico (slightly en-
larged). Fig. 2. From Lamarck’s original figure, 1816, Tableau Encyclopédique et Méthodique, Atlas 3,
pl. 422, fig. 3, Liste, p. 5.

A nomen nudum
Cymatium (Ranularia) mohorteri ‘Verrill’ Salisbury 1954, Zoological Record 89, sec. 9, p. 70 [nomen nudum].

This species was described in mimeographed form only.
ae he ag

The Genus Bailya M. Smith

In 1944 (Panamic Marine Shells, p. 22), Maxwell Smith instituted the genus Bailya
as anew genus in the family Cymatiidae. He included in this genus 7?7iton parvus C.B.
Adams. Unfortunately he had overlooked a previous publication by Pilsbry and Vanatta
(1904) in which 7. parvus C. B. Adams, on the basis of its soft anatomy, was considered
to be a species in T'ritonidea, subgenus Caducifer, which is in the Family Buccinidae.

This in no way invalidates Bailya, as the type species is Triton anomalus Hinds, an
Eastern Pacific species. Until the soft anatomy of this species is examined, the exact
position of Bailya will remain unknown.

244 JOHNSONIA, No. 36 References

REFERENCES

Bayer, Ch. 1933, Catalogue of the Cymatiidae in ’sRijks Museum van Natuurlijke Historie. Zoologische
Mededeelingen ’sRijks Museum van Natuurlijke Historie, Leiden, 16, pp. 33-59.

Belletante, H. 1954, Catalogue des Cymatiidae du Muséum National d’Histoire Naturelle de Paris. Journal
de Conchyliologie, 94, pp. 70-84; 85-96.

Carcelles, A. 1944, Catalogo de los Moluscos Marinos de Puerto Quequén. Revista del Museo de La Plata
(N.S.) Zoology, 3, pp. 233-309.

Dall, W.H. 1889, A Preliminary Catalogue of the Shell-Bearing Marine Mollusks of the Southeastern Coast
of the United States. Bull. United States National Museum, no. 37, pp. 1-221, 74 plates.

Dall, W.H. 1904, An Historical and Systematic Review of the Frog-Shells and Tritons. Smithsonian Mis-
cellaneous Collections, 47, pp. 114-144.

Emerson, W. K. and E. L. Puffer 1953, A Catalogue of the Molluscan Genus Distorsio. Proc. Biological
Society of Washington, 66, pp. 93-106.

Lange de Morretes, F. 1949, Ensaio de Catalogo dos Moluscos do Brasil. Arquivos do Museu Paranaense, 7,
pp. 1-216.

Lebour, M. V. 1945, The Eggs and Larvae of Some Prosobranchs from Bermuda. Proceedings Zoological
Society London, 114, pp. 462-489.

Locard, A. 1897, Expéditions Scientifiques du Travailleur et du Talisman, Paris, pp. 1-516, 22 plates.

Nicklés, M. 1950, Mollusques Testacés Marins de la Céte Occidentale d’Afrique. Manuels Ouest-Africains,
Paris, 2, pp. 1-269, 459 text figures.

Peile, A.J. 1926, The Mollusca of Bermuda. Proceedings Malacological Society London, 17, pp. 71-98.

Pilsbry, H. A. and E. G. Vanatta 1904, On Certain Rhachiglossate Gastropoda Eliminated from the Aquilli-
dae. Proc. Academy Natural Sciences Philadelphia, pp. 592-595.

Pilsbry, H.A. 1922, Revision of W.M.Gabb’s Tertiary Mollusca of Santo Domingo. Proc. Academy of Nat-
ural Sciences Philadelphia, 73, p. 357.

Pilsbry, H. A. and H. N. Lowe 1931. West Mexican and Central American Mollusks Collected by H.N.
Lowe. Proceedings Academy Natural Sciences Philadelphia, 84, pp. 33-144.

Puffer, E.L. 1953, Distorsio reticulata vs. Distorsio clathrata in the West Indies. Proc. of the Biological
Society of Washington, 66, pp. 109-124.

Smith, Maxwell 1944, Panamic Marine Shells. Winter Park, Florida, pp. 1-70, 53 plates.

Smith, Maxwell 1948, Triton, Helmet and Harp Shells. Winter Park, Florida, pp. 1-57, 16 plates.
Thiele, J. 1929, Handbuch der systematischen Weichtierkunde, 1, pp. 281-283.

Tryon, G.W. 1880, Manual of Conchology (1) 3, pp. 5-35, pls. 1-17.

von Ihering, H. 1907, Les Mollusques Fossiles du Tertiaire et du Crétacé Supérieur de ]’Argentine. Anales
del Museo Nacional de Buenos Aires (3) 7, 611 pages, 18 plates.

Watson, R.B. 1886, Voyage of H.M.S. Challenger, 15, pp. 389-396.
Wenz, W. 1941, Handbuch der Paldozoologie, 6, pt. 1, pp. 1056-1071.

Winckworth, R. 1945, The types of the Boltenian Genera. Proc. Malacological Society London, 26, pp.
136-148.

Woodring, Wendell P., 1928, Miocene Mollusks from Bowden, Jamaica, Part II. Carnegie Institution of
Washington, Publication no. 385, pp. 297-301.

pe
MUS. COMP. 7001

LIBRARY
JOHNSONIA MAY ~ 8 1958
pt Pabbahed shy salad
THe DEPARTMENT OF MOLLUSKS UNARERSITY

Museum of Comparative Zodlogy, Harvard University
Cambridge, Massachusetts

MAY 8, 1958 PHASIANELLIDAE VOL..335 NOL3T

THE FAMILY PHASIANELLIDAE IN THE WESTERN ATLANTIC
BY
RozpeErt ROBERTSON

The Phasianellidae is a family of gastropods found in shallow water in most tropical
and temperate seas. The shells are often small and are usually conspicuously colored and
patterned. The largest species live in the Australasian region.

The oldest fossil] phasianellids are found in European Paleocene deposits (Cossmann
1918). Reports of Cretaceous species are dubious. Like most families of marine mollusks
which differentiated in early Tertiary times, the group probably arose in Europe. The
European Eocene (Tethyan) marine mollusk fauna not only was in large part ancestral
to the modern Indo-Pacific fauna (see in particular Davies 1984) but also to the modern
Caribbean and Panamic faunas. The Phasianellidae were derived from a trochacean stock
and from Europe they spread around the world by way of the Tethyan Sea. Numerous
late Tertiary species have been described.

Plate 136. Phasianella australis Gmelin. Semaphore, near Port Adelaide, South Australia (1.5x). The type
species of Phasianella Lamarck.

245

246 JOHNSONIA, No. 37 Phasianellidae

The shell is entirely porcellaneous; in the most closely related families, the Turbinidae
and Trochidae, it is invariably nacreous within. Both in the Phasianellidae and the Tur-
binidae the operculum is calcareous and usually paucispiral, while in the Trochidae it is
always multispiral and corneous. The Phasianellidae differs from the Turbinidae and
Trochidae also in that the shells are usually smooth and bulimoid in shape rather than tur-
binate or trochoid. All three families belong in the superfamily Trochacea, which, in turn,
is grouped in the order Archaeogastropoda, the most primitive living gastropods. Most
families in this order have a rhipidoglossate radula, that is, have numerous marginal
teeth (the only exception is in the superfamily Patellacea—the true limpets). Primitively,
archaeogastropods have a pair of ctenidia (‘‘gills’’), but the Trochacea is one of the more
advanced groups in which one ctenidium (the one on the right side) has been lost as a
result of the long term effects of asymmetrical coiling.

Phasianellid radulae have been studied by 'Troschel (1878), Pilsbry (1888), Torr (1914),
Thiele (1924, 1929), Kuroda and Habe (1954) and Habe (1956). During the present
study the radulae of twenty species in the family were examined. Nine of these are type
species of supraspecific taxa. Material was not available of the types of the two remaining
groups based on Recent species (Pel/av Finlay and Gabrielona Iredale). All six Western
Atlantic species were examined.

There are four main kinds of phasianellid radulae. These correlate fairly well with
shell form and are here considered indicative of natural groups. Only two of these groups
live in the Western Atlantic.

In Phasianella Lamarck the central tooth of the radula is needlelike; often it is absent
along parts of the length of a ribbon. There are five pairs of laterals and each has an
elongate longitudinal attachment. Each transverse row of laterals is fairly straight (Plate
138, fig. 1). The complicated way in which the inner marginals fit into one another is
well described and figured by Torr (1914). This group lives from the Red Sea to New
Caledonia, north to Japan and south to Tasmania.

Plate 137. Fig. 1. Tricolia pullus Linné. St. Lunaire, near St. Malo, Ile et Vilaine, France (13.5x). The
type species of Tricolia Risso. Figs. 2-3. Gabrielona nepeanensis Gatliff and Gabriel. Point Nepean and
Flinders, Victoria, Australia (15x). The type species of Gabrielona Iredale. From Gatliff and Gabriel 1908,
pl. 21, figs. 9-10.

Western Atlantic JOHNSONIA, No. 37 ee

In Gabrielona the central tooth is large, with an entire cusp, and its attachment, wee,
the ribbon is neatly fitted between the innermost pair of laterals. There are five pairs -of
laterals and the transverse rows are fairly straight. The second marginal is large (Plate (, Cf
138, fig. 2). This group is little known and the description and figure are based on a°
West Indian species which is here referred to this genus on the basis of shell characters.
The type species, Phasianella nepeanensis Gatliff and Gabriel, is Australian.

In the third group, which includes Hiloa Pilsbry, Hotricolia Kuroda and Habe and
probably Pellax, free laterals are reduced in number to two or three pairs, each with a
‘hood.’ There is a cusped pseudocentral tooth made up of the reduced central which is
fused with the innermost pair of laterals. The median tooth of Hotricolia is considered a
true central by Kuroda and Habe (1954) and the three innermost pairs of marginals are
incorrectly considered laterals. Through comparisons with Hiloa it seems almost certain
that the median tooth of Kotricolia is a pseudocentral rather than a true central. Thiele’s
description (1929) of the radula of Hulithidium Pilsbry is based incorrectly on a species
in this group, which is known at present only from Hawaii, Australia and Japan.

In the fourth and largest group, T'ricolia Risso, s.s., the central tooth is large, mem-
branous, without cusps and variable in shape, even along the length of a single radula.
It is usually oval or rectangular. In all species examined there is a structure without
cusps, most easily seen through the ribbon, which lies between the outermost longitudi-
nal row of laterals and the innermost row of marginals, usually between the transverse
rows. The name lateromarginal plate is here suggested for it. Its function is problem-
atical and it appears not to have been previously described. The transverse rows of lat-
erals are M-shaped. All six European, African and western Indo-Pacific species so far
studied have five pairs of laterals (Plate 138, fig. 3). All the American species have four.’
In the three Eastern Pacific species of T?vicolia so far studied the innermost laterals are
elongate, winged laterally at the anterior end and lie close together beneath the central,
with the posterior part of the cusp projecting slightly over the central (Plate 138, figs.
4, 6). The cusps of the laterals in the two species figured are lobed, but in 7°. lurida
(Dall)’ and all five Western Atlantic species they are strongly dentate (Plate 138, fig. 5).
The innermost laterals of the Western Atlantic species differ in shape from the three
Eastern Pacific species in that they are less elongate and are not winged.

Reduction of the central tooth in this family has occurred in three ways in different
groups. In Phasianella this tooth has become needlelike or is often absent. In Hiloa
and related groups it has fused with the innermost pair of laterals. In T'ricolia, s.s., it
has become broad and membranous and is never cusped. In all three cases it seems to
be nonfunctional.

The marginal teeth are fairly similar in all four groups. They vary little in all trocha-
ceans. Juveniles seem to have fewer marginals than adults.

There are two main conclusions of phylogenetic significance to be drawn from this
brief discussion of phasianellid radulae. One is that within the family there is great diver-
sity in the structure of the radula. The other is that in many cases closely related species

1 Pilsbry (1888, pp. 163-164) implied that there is no phylogenetic significance in the number of laterals.
He ascribed the loss of the outermost pair to inefficiency of minute teeth in small species. This view is no
longer tenable as most of the American species are no smaller than the Paleotropical species.

> This name may not be applied correctly to this species. The holotype is apparently not the same species
as the one for which this name is often used. See Woodring (1957).

JOHNSONIA, No. 37 Phasianellidae

ibs)
i
Lo 6]

Plate 138. Radulae. Fig. 1. Phasianella australis Gmelin. South Australia (47x). Fig. 2. Gabrielona brevis
d’Orbigny. Antigua, Lesser Antilles (1370x). Fig. 3. Tricolia pullus Linné. St. Lunaire, France (280x).
Fig. 4. Tricolia compta Gould. Santa Barbara, California (280x). Fig. 5. Tricolia affinis affinis C. B. Adams.
Great Abaco Id., Bahama Islands (280x). Fig. 6. Tricolia variegata Carpenter. Bahia Magdalena, Baja
California, Mexico (280x).

Western Atlantic JOHNSONIA, No. 37 249

appear to live in the same region. For example, all the Western Atlantic species of
Tricolia appear to be more closely related to one another than to species elsewhere. This
is surprising and unusual, for as has often been pointed out, closely related species are
usually allopatric.

Phasianella jaws consist of two plates weakly joined at the dorsal margin and placed
directly above the anterior end of the radula. They probably serve to open the mouth
and provide a protective surface against which the radular teeth may rub. The radula
slides over the odontophore and abuts anteriorly against the jaws.

Pilsbry (1888, p. 162) describes the jaws in the whole family as ‘‘rhomboidal, covered
with imbricating scales.’ He illustrates (pl. 60, fig. 69) the jaws of T'ricolia fordiana
(Pilsbry) from Singapore. However, the jaws of Phasianella differ from those of T'ricolia.
Risbee has shown (1940, p. 283) that in P. variegata Lamarck [=P. rubens Lamarck]
they are not scaly. In this species the anterior edge of each plate is slightly thickened
and the outer surface is minutely reticulate. The jaws of this species are figured on Plate
139, figs. 1-2. In P. australis (Gmelin) they are much larger, corneous and fibrous and
vary considerably in thickness from specimen to specimen; some are thin and transparent
while others are much thicker and amber-colored. The differences are probably due in
part to age. The anterior edge of each plate is greatly thickened. The jaws of Phasianella
are externally convex and smooth with a large posteroventral gape between them, while
in Tricolia the plates are more or less flat and are at least occasionally scaly. Jaws may
prove to be an important taxonomic character in this family, but they are fairly variable
and difficult to study.

The epipodium in this family usually bears three pairs of cirri which presumably are
tactile for the epipodium is innervated, as in all rhipidoglossates (Pelseneer 1888). Both
the cirri and the tentacles bear fine immobile hairs. In 7vicolia there is usually a pair of
cervical lobes, also part of the epipodium. These have been described and figured for 7°.
pullus (Linné) by Forbes and Hanley (1849-50), by Clark (1855) and by Jeffreys (1865).
The left lobe is described by Pelseneer (1899) as being wider than the right, and Forbes
and Hanley, Clark, and Jeffreys all show that there is asymmetry in other features as
well, particularly in the amount of lobing. In 7°. bella (M. Smith) there is no lobe on
the right side; on the left it is pedunculate and digitate and serves as a sensory structure
which prevents particles in suspension in the ciliary current from entering the mantle
cavity (personal observations, Bimini, Bahama Islands). According to Pelseneer (1899)
the two lobes in 7. pudlus function like siphons, the one on the left being sensory, direct-
ing the ciliary current into the mantle cavity. Thus there is functional significance in the
asymmetry. Cervical lobes are lacking in Phasianella. However, in this genus there are
denticulate frontal lobes attached near the base of the tentacles. These are lacking in
Tricolia, as has been pointed out by Pilsbry (1888, pp. 164, 167). In all probability they
are not of epipodial origin, but are presumably sensory. Illustrations showing these
structures in Phasianella are given by Quoy and Gaimard (1832-85), Kiener (1847) and
Risbee (1940).

Fretter (1955) has discovered that there are two ‘‘shell’’ (columellar) muscles in T'ricolia
pullus, the right one being somewhat the larger. This is surprising for in all other ‘Troch-
acea that have been studied there is only one. Paired columellar muscles are known
in some of the Neritidae which have evolved toward a crepidula-like shell form, but it
seems unlikely that this indicates any close relationship between the two families as sug-

250 JOHNSONIA, No. 37 Phasianellidae

gested by Fretter. Whether or not these paired muscles are characteristic of the entire
family Phasianellidae is not known and should be investigated.

In both Phasianella and Tricolia the under side of the foot is divided by a median lon-
gitudinal furrow. The waves of progression are direct (posteroanterior) and ditaxic.

The microscopic structure of the eye of T7icolia pullus has been studied by Pelseneer
(1891) and the nervous system of Phasianella rubens by Risbec (1940).

According to Fretter (1955) the gut of 7. pudlus is similar to that in trochids except
that the two typhlosoles (folds) on the wall of the stomach continue along the length of
the intestine and rectum to the anus which is, as a result, functionally divided into two
openings. The feces are cylindrical rods, each with a longitudinal groove, and they are
extruded from one of the anal openings. The other opening probably acts as a siphon
allowing water to be expelled from the gut when the animal retracts into the shell.

As in all Trochacea the sexes are separate and there is no penis in the male. The sexes
can be distinguished (see Fretter 1955) by the appearance of the gonad and the size of
the urinogenital papilla (which is larger in the female). According to Lebour (1937) the
eggs of Tricolia pullus are shed singly into the sea and the veliger stage is short.

The shells in this family are usually bulimoid in shape and smooth except for fine spiral
sculpture. There are a few species of T'’ricolia, however, which have strong spiral cords,
such as 7". bella in the Western Atlantic and 7". bicarinata (Dunker) of South Africa.
The shell of Gabrielona is globose; a few species of T'’ricolia are more or less similar in
shape. Phasianella is much larger and usually higher spired than T’ricolia. The outer lip
of Tricolia is prosocline, that is, it grows farther forward at the suture than at the base,
and so is inclined relative to the axis. This is less marked in Gabrielona and Phasianella.
Periostracum is entirely lacking in the whole family.

The paucispiral, caleareous operculum is externally convex and more or less smooth in
Phasianella and Tricolia (Plate 139, figs. 6-8). In Gabrielona, on the other hand, it is
spirally ridged externally (Plate 139, figs. 8-4). Cossmann (1918) has pointed out that
in Phasianella there is a small parietal lamella within the aperture against which the oper-
culum pivots. This is not present in Gabrielona or Tricolia.

The fact that both primitive and juvenile turbinids often have a perforate operculum
has recently been discussed (Robertson 1957). Significantly, the juvenile operculum of
Tricolia bella is found to be perforate (Plate 139, fig. 5). This is further indication of the
fact that the Phasianellidae and Turbinidae are closely related.

There is but a single criterion by which the Phasianellidae are distinguished from the
Turbinidae: lack of nacre. According to Béoggild (1930) the microscopic structure of the
entire shell of Phasianella differs considerably from that of turbinids. The form of the
shell and the radula is often but not invariably diagnostic. There is also but a single cri-
terion by which these two families are distinguished from the Trochidae: the calcareous
operculum. The three families are nevertheless probably more or less natural groups.

In view of the fact that there are numerous differences between Phasianella and the
other groups which are now placed in the Phasianellidae, two subfamilies are here recog-
nized: the Phasianellinae, comprising the genus Phasianella, and the Tricoliinae, com-
prising the genera Gabrielona and Tricolia. The distinguishing characters are given below.
Very probably these two subfamilies were derived separately from the turbinid stock and
the similarities are due in part to convergence. The two groups are given subfamily rather
than family rank because the family Phasianellidae itself is at best hardly distinguishable

Western Atlantic JOHNSONIA, No. 37 251

from the Turbinidae. Thus the Phasianellidae may be considered diphyletic in origin.

The classifications of this family by Cossmann (1918) and Wenz (1938) overemphasize
differences which do not prove consistent. The phylogenetic series of species leading to
Recent forms suggested by Cossmann are greatly oversimplified. In some cases the re-
lationships he suggests are clearly erroneous, especially in the case of some of the West
Indian species.

The Eastern Pacific genus Prisogaster Mérch was thought by Thiele (1924, 1929) to
be intermediate in character between the Turbininae (='Turbinidae) and Phasianellinae

Plate 139. Figs. 1-2. Jaws of Phasianella rubens Lamarck. Caloundra, Queensland, Australia. Fig. 1, Oblique
posteroventral view. Fig. 2. Posteroventral view (both 38x). Figs. 3-4. Operculum of Gabrielona brevis
d’Orbigny. Antigua, Lesser Antilles. Fig. 3. External view. Fig. 4. Diagrammatic cross section (both
42x). Fig. 5. Juvenile specimen of Tricolia bella M. Smith showing perforate operculum. Pigeon Cays, An-
dros Id., Bahama Islands (50x). Figs. 6-8. Operculum of T'icolia affinis affinis C. B. Adams. Great Abaco
Id., Bahama Islands. Fig. 6. External view. Fig. 7. Diagrammatic cross section. Fig. 8. Internal view
(all 23x).

<9 JOHNSONIA, No. 37 Phasianellidae

L0=

(= Phasianellidae) and he grouped it with the latter. The cusp of the wide central tooth
of the radula in this genus is reduced and this suggested to Thiele affinity with T’ricolia,
s.s., in which there is no cusp at all. As the shell is nacreous within and turbinate in
form, and the radula is not closely similar to that of any phasianellid, the genus is here
referred back to the Turbinidae.

The genus Pseudophasianus Cossmann (1918), known only from Upper Eocene de-
posits in Europe, in all probability does not belong in the family. Azzyella Cossmann
(1889), based on a species from Eocene deposits near Paris, is often accorded generic
rank because of the spiral sculpture, but it is here ranked as a subgenus of T'ricolia for,
as mentioned above, there are Recent species which also have such spiral cords.

Some of the many supraspecific taxa in this family based on Recent species may be
given subgeneric rank under T'ricolia. The genus Tricolia, as here recognized, comprises
the third and fourth groups mentioned above in the discussion of radulae. Despite di-
vergence in structure of the radula there are few shell characters by which species in the
two radular groups may be separated. Species in the third group may be placed in Hiloa,
Eotricolia and possibly Pellax although differences between them are relatively slight.
The shells in this group are fairly thin, the suture is rather deeply impressed, the body
whorl is inflated, and the edge of the outer lip is sometimes everted. In the fourth group,
Tricolia, s.s., the shell is fairly thick and the body whorl is less inflated.

Most phasianellid species are remarkably variable, both in shell-form and in coloration.
One of the most important taxonomic characters is the color pattern. This is usually
fairly constant within a species. The shape of the shell is also moderately constant. Much
of the variability, particularly of color, is undoubtedly connected with the variety of sub-
strates selected (and hence food), as in Turbo (Ino 1955).

Despite the fact that Quoy and Gaimard report (1834) that Phasianella is attracted to
meat and that Clark (1855) invariably found foraminiferan tests in the stomach of 7'77-
colia pullus, this family, like most archaeogastropods, is predominantly herbivorous. On
the British coast 7°. pudlus lives in the lower intertidal and Laminarian zones, particularly
on the algae Chondrus, Ceramium, Rhodophyllis and Rhodymenia, and feeds on diatoms
and detritus as well as the surface tissues of these red algae (Jeffreys 1865; Fretter 1955).

The West Indian species of T'’ricolia live on both marine algae and grasses; a few live
among rocks, corals and even on sand, but these probably also graze on algae which are
present as incrusting films. 7" thalassicola Robertson has been dredged alive off Florida
in 85 fathoms; Forbes (1844) reports that 7". pullus lives as deep as 80 fathoms in the
Aegean Sea. Most species in this family are, however, confined to shallower water where
plant life is more abundant. Empty shells have been dredged in great depths in the
Caribbean, but these are of no significance as such small shells can easily be transported
in a variety of ways.

As far as is now known, T'ricolia affinis pterocladica Robertson lives only on the red
alga Pterocladia, and 7’. thalassicola predominantly on Turtle Grass (Thalassia testudt-
num Wonig). This is discussed further below. Attention is here drawn to a study by
Ostenfeld (1927) in which the distribution of Thalassia is mapped. This marine flowering
plant may determine the distribution of those herbivorous snails which live on it.

Tricola and Gabrielona are frequently drilled by predaceous gastropods (Plate 142,
figs. 1, 3). More often still the outer lip is broken back by fish, but usually such damage
is repaired by the animal. Brasilian specimens have been collected in the digestive tracts

Western Atlantic JOHNSONIA, No. 37 253

of sea hares (Aplysia) and of starfish (Haas 1953). Empty 7'ricolia shells, many of them
drilled, are remarkably abundant in sands throughout the West Indies and southern
Florida. Bleached and worn protoconchs are particularly in evidence. Surprisingly
enough, however, the genus is absent throughout most of the Gulf of Mexico, occurring
only sporadically on the Texan and Mexican coasts and off western Florida. It may also
be absent from Bermuda (see Notes).

Some of the syntypes (cotypes) of the five species of T’ricolia described from Jamaica
by C. B. Adams are mixtures of several species. This has been responsible for much con-
fusion as his specimens were widely disseminated. He described the species poorly and
some of the descriptions are composite.

Reeve (1862) and Sowerby (1884) incorrectly synonymized most of the West Indian
species which had been named at the time with T'ricolia pullus of Kurope. Records of
this species from the West Indies are thus erroneous, having been based on other species.

Pilsbry (1888) reviewed the family and laid the foundation for future work. Few tax-
onomic studies have been published on the Western Atlantic species since his excellent
monograph.

Strong (1928) studied the Eastern Pacific species of T’7ricolia. He recognizes 11 species,
3 of which he did not see. The genus Gabrielona appears not to be represented. As
discussed above, the radulae of the Eastern Pacific species of T'ricolia so far studied differ
consistently from those of the Western Atlantic. There are no clear-cut analogues in the
two oceans. The shells of 7°. cyclostoma (Carpenter) and 7’. thalassicola do, however, re-
semble one another, and several of the Eastern Pacific species have oblique spiral lines
of color similar to those in some of the Western Atlantic forms.

The oldest fossil American phasianellid from the Atlantic seaboard was recently
named T'ricolia calypta by Woodring (1957). It is from early Oligocene (or possibly late
Eocene) deposits of the Bohio (7) Formation, Gatun Lake area, Panama Canal Zone.
An early Miocene species was named Lacuna precursor by Dall (1915). It is from the
Tampa Limestone, Florida, and, as recognized by Mansfield (1937), it also is a Ticolia.
Other Miocene species so far described are: Lacuna punctata Gabb (1873) from the Cer-
eado Formation, Dominican Republic’; Phasianella *‘doubtful species’’ from Trinidad
(Guppy and Dall 1896); Phasianella mollis Olsson (1922) from the Gatun Formation,
Costa Rieca*; Tricolia (Eulithidium) hadra Woodring (1928) from the Bowden Forma-
tion, Jamaica (a Gabrielona’?); Tricolia probrevis Gardner (the holotype is juvenile),
Didianema ? waltonia Gardner (a Gabrielona?) and Tricolia affinis chipolana Gardner
(1947), all from the Alum Bluff Group, Florida, and T'ricolia ¢ syntoma Woodring
(1957) of the Gatun Formation, Panama (a Gabrielona?). \With the exception of the
three species which may be referred to Gabrielona all these Miocene forms are similar to
T.. affinis (C. B. Adams), a highly variable Recent species. Pliocene fossils have all been
referred to Recent species. It should be stated that the remarks made here about these

1 As there was a prior Tricolia punctata, Woodring (1928) renamed this 7’. (T'ricolia) affinis gabbi.

?This was synonymized by Woodring (1928) with 7’. umbilicata (d’Orbigny), a homonym, as shown in the
synonymies below, which has been applied to two Recent species: 7’. affinis and T. thalassicola. T. mollis may
be a synonym of 7’. qffinis, but the status of the name will remain in doubt until the type is examined. This
type should be in the Paleontological Research Institution, Ithaca, New York, but cannot at present be lo-
eated. Aguayo (1945) has suggested using Olsson’s name in place of 7’. wmbilicala,

954 JOHNSONIA, No. 37 Phasianellidae

fossils are made solely on the basis of descriptions and illustrations in the literature, not
on specimens. Hence they are especially open to question.

Finlay (1926) and Cotton and Godfrey (1938) use the name Eutropiidae (Eutropiinae
H. and A. Adams 1854) for the Phasianellidae. This is inadmissible because the name is
based on an objective junior synonym of Phasianella. The name Phasianellidae is already
well established. Cotton himself does not use the name /utropia.

ACKNOWLEDGMENTS

Without the help and cooperation of many individuals this work could not have been
completed. Approximately 20,000 Western Atlantic specimens were examined during
the study. Well over half of these were sent on loan. ‘To Dr. William J. Clench and to
Dr. Ruth D. Turner, under whose direction the work was carried out, I am greatly in-
debted. All the facilities of the Mollusk Department in the Museum of Comparative
Zodlogy (MCZ) were kindly placed at my disposal. To Dr. Myra Keen, Stanford Uni-
versity (LSJU), under whom the work was started, 1 am also deeply grateful, and to
those who so kindly sent material on loan and in many cases gave important informa-
tion as well. I would particularly hke to mention Dr. R. Tucker Abbott, Academy of
Natural Sciences of Philadelphia (ANSP); Dr. Fritz Haas, Chicago Natural History
Museum (CNHM);: Dr. Leo G. Hertlein, California Academy of Sciences (CAS): Mr.
Thomas L. McGinty, Boynton Beach, Florida; Dr. Donald F. McMichael, Australian
Museum, Sydney; Dr. Harald A. Rehder, United States National Museum (USNM),
and Mrs. Germaine L. Warmke, Mayagiiez, Puerto Rico. The following also very kindly
sent material on loan: Mr. R. M. DeWitt, Florida State Museum (FSM); Mr. George
F. Kline, Madison, New Jersey; Mr. Donald R. Moore, Ocean Springs, Mississippi: Dr.
and Mrs. David Schmidt, Sarona, Wisconsin: Dr. Henry van der Schalie, Museum of
Zoology, University of Michigan (U of M): Dr. Gilbert L. Voss, University of Miami
Marine Laboratory (ML), and Mr. Frederick V. Weir, American Museum of Natural
History (AMNH).

I am under obligation to the late Guy L. Wilkins and to I. C. J. Galbraith, both of
the British Museum (Natural History), for the photographs of the types illustrated on
Plate 142.

Dr. William Randolph Taylor, Department of Botany, University of Michigan, iden-
tified an alga, and Dr. Wendell P. Woodring, U.S. Geological Survey and Dr. K.V.W.
Palmer, Paleontological Research Institution, kindly answered several queries.

Some manuscript notes made by the late Dr. Henry A. Pilsbry on the Western At-
lantic species of T'ricolia were kindly sent on loan toward the end of the study.

Family PHASIANELLIDAE

Subfamily PHAsSIANELLINAE

Adult shell 1 to 10 em. in length, oval, smooth and polished. Spire high, produced at
an angle of from 45° to 55°. Whorls evenly rounded. There is a callus on the parietal
region but no umbilicus. The color pattern is made up in part of spiral lines or bands.
The operculum, which is externally convex and more or less smooth, pivots against a
weak axial lamella on the columella, well within the aperture. The aperture is oval and
the outer lip fairly thin and weakly prosocline.

Western Atlantic JOHNSONIA, No. 37 255

The central tooth of the radula, when present, is needlelike. Each of the five paired
laterals has an entire cusp and an elongate longitudinal attachment to the ribbon: the
transverse rows are fairly straight. The plates of the jaw are externally convex.

Frontal lobes are present on the head, but there are no cervical lobes.

Genus Phasianella Lamarck

Phasianella Lamarck 1804, Annales du Muséum National d’ Histoire Naturelle (Paris) 4, p. 295.
Phasianus Montfort 1810, Conchyliologie Systématique 2, p. 255 [emendation of Phasianella Lamarck |

non Phasianus Linné 1758 [Aves]. .
Bolina Rafinesque 1815, Analyse de la Nature, Palermo, p. 144 [new name for Phasianella Lamarck }.
Eutropia ‘Humphrey’ Swainson 1840, Treatise on Malacology, p. 21 [merely listed as a synonym of Phas-

ianella Lamarck |; ‘Humphrey’ H. and A. Adams 1854, Genera of Recent Mollusca, London, 1, pp. 389-390.
Orthopnoea Gistel and Bromme [1847] 1850, Handbuch der Naturgeschichte, p. 556; Gistel 1848, Natur-

geschichte des Thierreichs fiir héhere Schulen, pp. 169, 199 [new name for Phasianella Lamarck }.
Orthomesus Pilsbry 1888, Manual of Conchology (1) 10, pp. 163, 179 (type species, Phasianella variegata

Lamarck 1822, non P. variegata de Roissy 1805 [=P. rubens Lamarck 1822], by original designation).
Mimelenchus Iredale 1924, Proceedings of the Linnean Society of New South Wales 49, pp. 182, 232 (type

species, Phasianella ventricosa Swainson‘ 1822, by monotypy).

Type species: ‘‘faisan’’ [= Buccinum australe Gmelin 1791], by subsequent selection,
de Roissy 1805.

The characters given above for the subfamily apply to the genus.

Lamarck based his description of the genus on the ‘‘faisan’” but in the same paper de-
scribed two fossil species (now referred to Tvicolia) from the Paris Basin. Harris (1897)
has selected Phasianella turbinoides Lamarck 1804 as type. Woodring (1928) argued that
the type species is the ‘‘faisan”’ by original designation. Lamarck, however, did not use
the word ‘type’ when referring to this species. Fortunately de Roissy selected it as type
in 1805, as has been discussed recently by Woodring (1957). The selection made by
Harris is therefore invalid.

Neither the shell nor the radula of Orthomesus or Mimelenchus differs in any significant
way from Phasianella, s.s.

A figure of a radula published by Eberhard (1865) which was believed by 'Troschel
(1878) to be that of Phasianella australis has caused much confusion, for it was a T'7ico-
lia (probably T. pullus), not a Phasianella. Pilsbry named Orthomesus as a result of this
misapprehension. For a long time the major differences between the radulae of Phasi-
anella and Tricolia were thus not realized. Iredale (1924) and Thiele (1924) were the
first to point this out.

The oldest undoubted representatives of this genus appear to be from rocks of Miocene
age in Java and Victoria, Australia (Martin 1916; Crespin 1926).

The following synonymy is of the type species of the genus.

Phasianella australis Gmelin
Plate 136; Plate 138, fig. 1

Buccinum australe Gmelin 1791, Systema Naturae, ed. 13, p. 3490 (rivers of New Zealand) [based on Fa-
vanne de Montcervelle 1784, Catalogue Systématique et Raisonné ou Description du Magnifique Cabinet Ap-

1 See Iredale’s discussion (1924) of Swainson’s use of this name prior to Quoy and Gaimard (1834) for the
same species.

256 JOHNSONIA, No. 37 Phasianellidae

partenant ci-devant a M. le C[omte] de [La Tour d’Auvergne], p. 11, pl. 1, fig. 46, and Chemnitz 1786,
Conchylien-Cabinet (1) 9, pt. 2, pp. 38-40, pl. 120, figs. 1033-1034].

Helix phasianus Riding 1798, Museum Boltenianum, p. 108 [based on Chemnitz 1786, pl. 120, figs. 1033-
1034].

Phasianella variegata’ de Roissy 1805, Histoire Naturelle Générale et Particuliére des Mollusques 5, p. 331
(islands near New Holland [ Australia } ) [based on Chemnitz 1786, pl. 120, figs. 1033-1034].

Phasianus variegatus de Roissy. Montfort 1810, Conchyliologie Systématique 2, p. 255.

Bulimus phasianus Perry 1810, Areana, sign. Y, 7th page, pl. 43 (New Holland and Van Diemen’s Land
{Australia and Tasmania] ); Perry 1811, Conchology, or the Natural History of Shells, pl. 30, fig. 4.

Trochus phasianella ? Brookes 1815, An Introduction to the study of Conchology, p. 163, pl. 7, fig. 96.

Phasianella varia Lamarck 1816, Tableau Encyclopédique et Méthodique, Mollusques et Polypes Divers,
Liste, p. 10, pl. 449, figs. la, b, c. ;

Phasianella bulimoides Lamarck 1822, Histoire Naturelle des Animaux sans Vertébres 7, p. 52 (seas of New
Zealand and New Holland [Australia], common near Maria Id. [off eastern Tasmania] ) [based on Buccinum
australe Gmelin and the preceding].

Phasianella picta de Blainville 1825, Manuel de Malacologie, p. 439; 1827, pl. 37, figs. 5, 5a.

Phasianella tritonis ‘Chemnitz’ Anton 1839, Verzeichniss Conchylien, Halle, p. 60.

Phasianella lehmanni Menke 1843, Molluscorum Novae’ Hollandiae, p. 12.

Phasianella preissii Menke 1843,, Molluscorum Novae Hollandiae, p. 12.

Phasianella venusta Reeve 1848, Elements of Conchology 1, p. 132, pl. 12, fig. 58.

Phasianella decorata Chenu 1859, Manuel de Conchyliologie 1, pp. 342-343, fig. 2530.

Phasianella pulchella Tenison-Woods 1877, Papers and Proceedings and Report of the Royal Society of Tas-
mania for 1876, p. 141 (Long Bay, Tasmania) [juvenile]; non P. pulchella Récluz 1843.

Phasianella delicatula Tenison-Woods 1878, Papers and Proceedings and Report of the Royal Society of Tas-
mania for 1877, p. 38 [new name for P. pulchella Tenison-Woods 1877, non P. pulchella Récluz 1843].

Phasianella australis Gmelin var. subsanguinea Pilsbry 1888, Manual of Conchology (1) 10, p. 165, pl. 38,

fig. 52.

This species ranges from Western Australia east along the south coast to Victoria and
‘Tasmania. The type locality (rivers of New Zealand) is erroneous. For further informa-
tion see Cotton (19435).

Subfamily TricoLiinakE

Adult shell normally less than 1 cm. in length. Spire usually produced at an angle of
60° or more. Shape and sculpture various, but spiral grooves are often present, particu-
larly on the early whorls. Small species are perforate; large species are imperforate when
adult. Color pattern various. Aperture oval or semicircular. A parietal lamella is absent.
In Tricolia the operculum is similar to that of Phastanella; in Gabrielona it is naticoid.

The central tooth of the radula may have an entire cusp, may be reduced to a mem-
branous plate, or may be lacking altogether. In the latter case the innermost laterals
have fused to form a pseudocentral tooth and the free laterals are reduced in number to
two or three pairs.

There are cervical lobes but no frontal lobes on the head.

'This name invalidates Phasianella variegata Lamarck 1822 (Histoire Naturelle des Animaux sans Vertébres
7, p. 53). The species to which this name has been applied may take the name Phasianella rubens, introduced
by Lamarck on the same page for a different (possibly geographic) form of the same species. Iredale (1924)
and Cotton (1945) believe these are distinct species; this is here believed incorrect. If, however, it can be
shown that they are specifically distinct, the name Turbo lineolatus Wood (1828, Index Testaceologicus, Sup-
plement, p. 19, pl. 6, Turbo, fig. 26 [Phasianella lineolatus, p. 48|) is available instead of P. rubens for P.
variegata Lamarck.

a

Western Atlantic JOHNSONIA, No. 37 g

©

Fy
91

Woodring (1928) applied the name Tricoliidae to this group, separating it from the
Phasianellidae.

Genus Gabrielona Iredale

Gabrielona Iredale 1917, Proceedings Malacological Society of London 12, p. 327.
Gabrielana ‘Iredale’ Cossmann 1918, Revue Critique de Paléozoologie 22, p. 42 [error for Gabrielona|.

Type species: Phasianella nepeanensis Gatliff and Gabriel 1908, by monotypy.

Adult shell not more than 2.4 mm. in length, globose, wider than long. Spire low,
usually produced at an angle of about 110°. Shell smooth or with fine incised spiral
grooves. An umbilicus is present. Whorls inflated. Shell pink or red-brown and white.
Aperture semicircular with the outer lip weakly prosocline. The operculum spirally
ridged externally, thinnest near the center and thickest near the margin. The central
tooth of the radula is large and has an entire cusp. Five pairs of laterals are present, the
cusps of which are also entire.

In their original description of the type species of Gabrielona, Gatliff and Gabriel
(1908) mention a similarity to the type species of Chromotis H. and A. Adams. This
no doubt induced Thiele (1929) to synonymize Gabrielona tentatively with Chromotis.
However, Chromotis is a subjective synonym of T'ricolia based on a species with an ex-
ternally convex operculum. As stated by Iredale, the operculum of Gabrielona is nati-
coid and, in addition, there are marked differences in the shell by which this genus differs
from T'ricolia.

Phasianella brevis d’Orbigny is here referred to this genus because of the similarity of
the operculum, color pattern and shell form to that of the type species. Dall (1889a)
wrongly referred it to Hucosmia Carpenter [= Hulithidium]. As discussed in the intro-
duction, the radula of this species is of a distinct form.

Tricolia (Eulithidium) hadra Woodring (1928), Didianema ? waltonia Gardner (1947)
and T'ricolia ? syntoma Woodring (1957) may provisionally be referred also to this genus,
pending knowledge of the opercula of these Miocene forms.

Plate 140. Fig. 1. Gabrielona nepeanensis Gatliff and Gabriel. Ocean Beach, near Point Nepean, Victoria,
Australia (27x). Paratype, Australian Museum, no. C. 45057. Juvenile specimen with a slightly broken
outer lip. Figs. 2-3. Gabrielona brevis d’Orbigny. Fig. 2. Antigua, Lesser Antilles (28x). Fig. 3. Off Bahia
Honda, Pinar del Rio, Cuba (25.5x). Same specimen figured by Dall 1889, pl. 19, fig. 10b.

258 JOHNSONIA, No. 37 Gabrielona

Gabrielona nepeanensis Gatliff and Gabriel
Plate 137, figs. 2-3; Plate 140, fig. 1

Phasianella nepeanensis Gatliff and Gabriel 1908, Proceedings Royal Society of Victoria (a.s:) 21, p. 366,
pl. 21, figs. 9-10 [reproduced here] (Flinders, Western Port; Ocean Beach, near Point Nepean [Victoria,
Australia] ); Iredale 1917, Proceedings Malacological Society of London 12, p. 327.

Gabrielona nepeanensis Gatliff and Gabriel. Cotton 1945, Transactions Royal Society of South Australia

69, p. 165.

This, the type species of Gabrielona, is known only from the type locality in Victoria
and possibly from two localities in South Australia. It is likely that Cotton (1945) has
misidentified this species, for he states that his specimens are separated with difficulty
from ‘‘Pellax virgo.’’ The two species are easily distinguished, for they differ greatly in
shape and in the structure of the operculum.

Gabrielona brevis d’Orbigny
Plate 138, fig. 2; Plate 139, figs. 3-4; Plate 140, figs. 2-8; Plate 141; Plate 142, fig. 1

Phasianella brevis d’Orbigny 1842 [in] Ramon de la Sagra, Histoire de ]’Ile de Cuba, Mollusques 2, p. 79,
pl. 20, figs. 19-21 (Martinique and Cuba).

Phasianella brevissima Pilsbry 1888, Manual of Conchology (1) 10, p. 179 [new name for P. brevis d’Orbigny
1842, non P. brevis C. B. Adams 1850].

Phasianella (Eucosmia) brevis d’Orbigny. Dall 1889, Bulletin Museum of Comparative Zoédlogy 18, p. 351,
pl. 19, fig. 10b [ Eucosmia brevis on plate explanation].

Eucosmia brevis d’Orbigny. Cossmann 1918, Essais de Paléoconchologie Comparée, Paris, livr. 11, p. 162,

fig. 55.

Tricolia brevis d’Orbigny. Aguayo 1943, Revista de la Sociedad Malacolégica 1, pp. 39-40.

Description. Shell moderately thin, globose, inflated and reaching 2.4 mm. in length.
The color pattern is made up of light to dark brick-red irregular wavy stripes on white.
The columellar area is either white and demarcated from the surrounding light brick-red
by a crenulate margin, or there are more or less square white spots in a spiral series on
dark brick-red. The wavy stripes are widest below the suture and below the periphery.
There may be merely series of flames above and below the periphery on the initial whorls.
In one specimen the dark wavy stripes are so wide that they have fused to leave wavy
white lines on a dark brick-red ground color. Whorls 4, greatly inflated. Spire depressed,
set at an angle of 110°. Protoconch minute, depressed and white. Aperture pyriform-
ovate. Columellar callus fairly thick, margin raised slightly near the umbilicus. Umbil-
icus narrow. Suture slightly impressed. Post-nuclear whorls usually with evenly spaced
spiral striae; the striae gradually become weaker so that the body whorl is entirely
smooth. One specimen is entirely smooth and polished. Operculum (Plate 139, figs. 3-4)
naticoid, more or less flat, but ridged near the margin and thinnest at the center.

length width
2.4mm. 2.2mm. Antigua, Lesser Antilles

2.0 2.4 Off Bahia Honda, Cuba
s5 1.6 Antigua, Lesser Antilles
(juvenile) he 1.4 Habana, Cuba

Types. Three syntypes of Phasianella brevis d’Orbigny (one is illustrated here, Plate
142, fig. 1) are in the British Museum (Natural History), no. 54.10.4.282. Gray (1854)

Western Atlantic JOHNSONIA, No. 37 259

states that all the d’°Orbigny types from Cuba were sent to the Museum.’ They are from
Martinique, however, not Cuba as implied by the title of Gray’s list. The Cuban speci-
men(s) mentioned by d’Orbigny may thus have been lost, or were not regarded as types
and retained by d’Orbigny. The type locality is Martinique, Lesser Antilles.

Remarks. 'This species can easily be distinguished from other Western Atlantic phas-
ianellids because the shell is as wide as it is long, or almost so, and the operculum is
naticoid. The light to dark brick-red color is also characteristic; there are no spots.

Only thirteen specimens were available for this study. Eleven of them were collected
at Antigua by J. B. Henderson, Jr.; one was dredged dead in deep water off Bahia
Honda, Cuba, by the Blake, and one juvenile is from near Habana, Cuba. The Cuban
specimen from deep water differs from the Antiguan specimens in that it is smooth and
predominantly dark brick-red, with wavy white streaks, and the outer lip is thickened.
All the Antiguan specimens have spiral striae, particularly on the early whorls. They
are white, with wavy pale brick-red stripes. The outer lip is thin, even though some of
the specimens are larger than the one from Cuba.

One of the Antiguan specimens was collected alive in ‘‘deep, fine sand and shells,”’
apparently in shallow water.

Dall (1889a) identified the Cuban specimen from deep water with d’Orbigny’s species:
however, he misidentified the deep water form of T?7colia thalassicola with this species
also, as well as juvenile specimens of other West Indian species. The published ranges
of this species have been incorrect as a result.

Tricolia probrevis Gardner (1947) was implied to be a precursor of this species. How-
ever, it is a Juvenile Tvicolia, not a Gabrielona. The specimens with which it was com-
pared were probably misidentified.

Six worn and broken specimens in the United States National Museum (no. 150781)
from Ilha de Sao Sebastiaéo, Sio Paulo, Brasil, collected by H. von Ihering, may possi-
bly represent another Western Atlantic species of Gabrielona. What little remains of
the color pattern consists of irregular red-brown spots. The spire is higher than in G.
brevis, being produced at an angle of about 85°, and the umbilicus is more prominent.
The largest specimen is 1.9 mm. in length. Several of the specimens show traces of spiral
striae. The Brasilian record of **Hulithidium brevissimum (Pilsbry)’’ given by Lange de
Morretes (1949) may have been based on specimens from the same lot.

Plate 141. Gabrielona brevis d’Orbigny. Fig. 1. Antigua, Lesser Antilles. Fig. 2. Off Bahia Honda, Pinar
del Rio, Cuba. Same specimen figured by Dall 1889, pl. 19, fig. 10b (both 13x).

‘In the notes made by P. P. Carpenter while at the Museum in 1863-64 which have been published by
Palmer (1947) it is stated that these specimens are ‘‘a Glabella, very like the commoner C. S. Lucas sp.”
[Cape San Lucas, Baja California, Mexico]. Glabella Swainson is a marginellid; possibly Carpenter intended
to use this name for the group he shortly afterwards called Eucosmia in a paper describing new species from

Cape San Lucas.

260 JOHNSONIA, No. 37 Tricolia

Range. Cuba, Antigua and Martinique (d’Orbigny).

Specimens examined. Cua: Blake, station 21 [1877-78] (N. Lat. 23°2’; W. Long.
83°13’) in 287 fathoms (dead), off Bahia Honda, Pinar del Rio (USNM); Arenas de la
Chorrera, Habana (MCZ). Lesser ANTILLES: English Harbour, Antigua (USNM).

Genus Tricolia Risso

Tricolia Risso 1826, Histoire Naturelle de 1’Europe Méridionale, Paris, 4, p. 122.

Tricolea ‘‘Risso,’’ Thicolea “‘Leach MSS. 1819,’’ Tricolaea, all Gray 1847, Proceedings of the Zoological
Society of London 15, pp. 144, 218 {errors for Ticolia Risso].

Eudora ‘Leach’ Gray 1852, Synopsis of the Mollusca of Great Britain, pp. 147, 199 (type species, Eudora
varians ‘Leach’ Gray 1852 [ = Turbo pullus Linné 1758], by monotypy); non Eudora Péron and Lesueur 1810
[Coelenterata |, ete.

Chromotis H. and A. Adams 1863, Annals and Magazine of Natural History (3) 11, pp. 19-20 (type species,
Chromotis neritina Dunker { = Phasianella neritina Dunker 1846], by monotypy).

Eucosmia Carpenter 1864, Annals and Magazine of Natural History (3) 13, p. 475 (type species, Eucosmia
variegata Carpenter 1864, by subsequent selection, Pilsbry 1888); non Eucosmia Stephens 1831 [ Lepidoptera }.

Tricoliella Monterosato 1884, I] Naturalista Siciliano 3, p. 110 (type species, T’ricolia pullus Linné 1758,
by subsequent selection, Pilsbry 1888).

Steganomphalus Harris and Burrows 1891, Eocene & Oligocene Beds of the Paris Basin, London, pp. 78—
79, 112 [new name for Eudora ‘Leach’ Gray 1852, non Eudora Péron and Lesueur 1810, ete. ].

Eulithidium Pilsbry 1898, Nautilus 12, p. 60 [new name for Eucosmia Carpenter 1864, non Eucosmia Stephens
183i}.

Tricola Strong 1928, Proceedings California Academy of Sciences (4) 17, pp. 189, 191-193, 202 [error for
Tricolia Risso].

Usatricolia Habe 1956, Venus 19, pp. 95-96, 98 (type species, Phasianella compta Gould 1855, by monotypy).

Type species: Turbo pullus Linné 1758 [= Tricola pullus Linné], by subsequent se-
lection, Gray 1847 [misspelled ‘‘ 7'’ricolea’’ }.

Adult shell rarely over 1 cm. in length, globose, oval or elongate, usually smooth, but
a few species have strong spiral cords. Spire moderately elevated, usually produced at
an angle of from 60° to 80°, but very occasionally as low as 100° or as high as 40°. In-
itial whorls sometimes with strong spiral threads. Shell imperforate or rimately perforate.
Color pattern various, often consisting in part of oblique spiral lines. Aperture oval.
Outer lip strongly prosocline. Operculum white, externally convex and more or less
smooth, perforate when juvenile. Cervical lobes are present, but not frontal lobes. The
plates of the jaw are more or less flat.

The type species of Chromotis, from South Africa, is low-spired. However, the radula
differs in no observable way from that of T’ricolia pullus and the operculum has the same
general form as in all other species of T'ricolia. As there are several species transitional
in shell form, it seems unnecessary to maintain such monotypic subgenera.

Low-spired forms of T’ricolia (especially American species) have frequently been re-
ferred to Hulithidium. However, the spire of the type species (Plate 148, fig. 1) is no
lower than in T'ricolia, s.s. (Plate 137, fig. 1). The assemblage of species referred by Dall
(1908) to this subgenus is clearly heterogeneous. As discussed in the introduction, the
radulae of the American species of T'ricolia differ consistently from those from the Old
World in having four rather than five pairs of radular laterals. The name Hulithidium
could be applied subgenerically to the American species, but at the present time it is

Western Atlantic JOHNSONIA, No. 37 261

thought that this would be confusing for this name has long been applied incorrectly only
to low-spired species.

Usatricolia was separated from T'ricolia because the central tooth of the radula was
‘distinctly transversely narrower’’ and because there were four pairs of laterals rather
than five. There is great variability in the shape of the central tooth, even in a single
radula. The tooth figured by Habe is merely abraded. The radula of Usatricolia (Plate
138, fig. 4) is strikingly similar to that of Hulithidium (Plate 138, fig. 6) and the two
cannot possibly be considered even subgenerically distinct (Plate 148). Both have four
pairs of laterals.

Six subgenera may be recognized within the genus: T'ricolia, s.s., Aizyella Cossmann
(1889), Phasianochilus Cossmann (1918), Hiloa Pilsbry (1917), Pellaw Finlay (1926) and
Eotricolia Kuroda and Habe (1954).

Subgenus Tricolia

Shell fairly thick; suture never deeply impressed; whorls moderately inflated. Cen-
tral tooth of the radula not cusped, reduced to a membranous plate. Four or five pairs
of laterals are present, each with incised cusps. Rows of lateromarginal plates lie between
the marginals and laterals.

All the Western Atlantic species belong in this subgenus.

Varietal names are omitted in the following synonymy of the type species. For more
complete synonymies, which include some of the many varietal names, see Bucquoy,
Dautzenberg and Dollfus (1884) and Priolo (1953).

Tricolia pullus Linné
Plate 137, fig. 1; Plate 138, fig. 3

Turbo pullus’ Linné 1758, Systema Naturae, ed. 10, p. 761 (Mediterranean Sea); Hanley 1855, Ipsa Lin-
naei Conchylia, pp. 327-328.

Turbo pictus da Costa 1778, British Conchology, p. 103, pl. 8, figs. 1, 3 (coast of Cornwall, and Exmouth,
Devonshire; Mediterranean) [refers to the preceding].

Turbo flammeus von Salis 1793, Reisen in versch. Prov. Kénigreich Neapel, p. 377, pl. 8, fig. 11 (Naples,
Italy). Not seen.

Phasianella pullus Linné. Sowerby 1825, Catalogue of the Shells contained in the collection of the late Earl
of Tankerville, p. 56.

Tricolia pullus Linné. Risso 1826, Histoire Naturelle de 1’Europe Méridionale, Paris, 4, p. 123.

Tricolia punctata Risso 1826, Histoire Naturelle de 1’Europe Méridionale 4, p. 123 (coast of Provence).

Phasianella pullulus Anton 1839, Verzeichniss Conchylien, Halle, p. 60.

Phasianella pulchella Récluz 1843, Revue Zoologique par la Société Cuvierienne 6, pp. 10-11 (coasts of the
English Channel).

Phasianella tenuis Philippi 1844, Zeitschrift fiir Malakozoologie 1, p. 110 (Sicily); non P. tenuis Michaud
1829.

Eudora varians ‘Leach’ Gray 1852, Synopsis of the Mollusca of Great Britain, p. 200 [new name for Turbo
pullus Linné 1758].

Phasianella crassa Brusina 1864, Verhandlungen zoologisch-botanischen Gesellschaft in Wien 15, p. 23
(Dalmatia).

? Phasianella exigua Brusina 1864, Verhandlungen zoologisch-botanischen Gesellschaft in Wien 15, p. 24
(Dalmatia).

' The name Pullus was capitalized by Linné, indicating that he used it asa noun. See Cooke 1899, Journal
of Malacology 7, pp. 31-32.

262 JOHNSONIA, No. 37 Tricolia

? Tricoliella jolyi Monterosato 1889, Journal de Conchyliologie 37, p. 30 (Casablanca and near Algiers).
Eudora dubia Monterosato 1889, Journal de Conchyliologie 37, p. 31 (Casablanca).

This species ranges from the British Isles south to West Africa, west to the Azores
and east along the shores of the Mediterranean to the Black Sea.

4

Plate 142. Figs. 1-3. D’Orbigny types. Fig. 1. Phasianella brevis | = Gabrielona brevis]. Syntype, Martinique,
Lesser Antilles (6x). Fig. 2. Phasianella umbilicata [= Tricolia thalassicola Robertson]. Invalid syntype,
Cuba (4x). Fig. 3. Phasianella zebrina [= Tricolia tessellata Potiez and Michaud]. Syntype, Guadeloupe,
Lesser Antilles (6x). Figs. 4-5. Syntypes of Littorina adamsii Reeve [| = Tricolia bella M. Smith]. Jamaica
(both 6x). Photographs by courtesy of the British Museum (Natural History).

Tricolia affinis affinis C. B. Adams
Plate 138, fig. 5; Plate 139, figs. 6-8; Plate 143, figs. 1-2; Plate 145, fig. 1

Phasianella umbilicata d’Orbigny 1842 [in] Ramon de la Sagra, Histoire de 1’Ile de Cuba, Mollusques 2,
p. 77, pl. 19, figs. 32-34 (Martinique, Guadeloupe, Jamaica and Cuba) [in part only]; non Littorina umbilicata
d’Orbigny 1840 [a Tricolia].

Phasianella affinis C. B. Adams 1850, Contributions to Conchology, no. 4, p. 67 (Jamaica).

Phasianella concinna C. B. Adams 1850, Contributions to Conchology, no. 5, p. 69 (Jamaica).

Description. Shell fairly thin, inflated, elongate, reaching 8 mm. in length. The color
pattern consists of irregularly arranged minute brick-red or red dots each of which is
usually paired with a somewhat larger white spot. These are produced subsequent to
each red dot in spiral sequence. The ground color is light orange-brown or pink and is
partially transparent. Irregular wavy axial streaks are superimposed which are either red
or alternately light brick-red and white. The white streaks are often broken at the pe-
riphery. Occasionally the whole shell is suffused with rose. Whorls 5, evenly rounded.
Spire produced at an angle of from 60° to 63°. Protoconch minute, depressed and
colored white or pale pink. Aperture elongate-ovate. Columella with a thick white cal-
lus. Umbilicus fairly wide. Suture impressed. Post-nuclear whorls smooth except for
minute spiral striae. Operculum elongate, white and minutely striated exteriorly. The
striae are near and at right angles to the margin.

length width
7.5mm. 5.0mm. _ Lectotype of Phasianella affinis C. B. Adams

6.8 4.2 Norris Cut, near Miami, Florida
5.0 3.2 Aguadilla, Puerto Rico
(juvenile) 2.7 1.9 Big Pine Key, Florida

Types. 'The lectotype of Phasianella affinis C. B. Adams (Plate 143, fig. 2), selected
by Clench and Turner (1950, pl. 36, fig. 6), is in the Museum of Comparative Zodélogy,
no. 186020. Lectoparatypes from Jamaica are in the United States National Museum

Western Atlantic JOHNSONIA, No. 37 263

(no. 61764) and the Academy of Natural Sciences of Philadelphia (no. 199422). The
lectotype of Phasianella concinna C. B. Adams (Plate 1438, fig. 1), also selected by Clench
and Turner (1950, pl. 36, fig. 4), is in the United States National Museum, no. 54766.
See under 7’. thalassicola concerning Phasianella umbilicata d’Orbigny. The type locality
of this species is Jamaica.

Remarks. This is the largest and one of the most abundant forms of 7J'ricolia in the
West Indies. See Remarks on other species and subspecies of T'ricolia for characters by
which it may be distinguished. In the Florida Keys it lives predominantly on Turtle
Grass (Thalassia testudinum Konig); at Great Abaco, in the northern Bahama Islands,
it was collected alive by the writer both in deep crevices in dead coral and on Thalassia.

Range. The Florida Keys, the Bahama Islands, the Greater Antilles east to Saint
Martin, northern Lesser Antilles. Replaced in most of the remainder of the Lesser An-
tilles by 7°. affinis beawi, by T’. affinis cruenta on the continental shores of the Caribbean and
the coast of Brasil, and by 7". affinis pterocladica north of the vicinity of Miami, Florida.

Specimens examined. FLortoA: Miami Beach (FSM); Fisher Id., Miami (T. McGinty);
Norris Cut (MCZ); Key Largo (MCZ; ANSP; ML; CNHM;; D. Moore); Teatable
Key (FSM; T. McGinty): Lower Matecumbe Key (USNM); Little Duck Key (T.
McGinty); West Sister Key, off Vaca Key (FSM); Bahia Honda Key (T. McGinty;

Plate 143. C.B. Adams types. Fig. 1. Phasianella concinna [= Tricolia affinis affinis C.B. Adams]. Lectotype,
Jamaica (4x). Fig. 2. Phasianella affinis | = Tricolia affinis affinis|. Lectotype, Jamaica (8x). Fig. 8. Phas-
ianella brevis | = Tricolia adamsi Philippi]. Holotype, Jamaica (16.5x). Fig. 4. Phasianella brevis {| = Tricolia
adamsi Philippi]. Invalid neotype, St. Thomas, Virgin Islands (15x). Fig. 5. Turbo (?) pulchellus | = Tri-
colia bella M. Smith]. Lectotype, Jamaica (10x). Fig. 6. Phasianella tesselata [= Tricolia tessellata Potiez
and Michaud]. Lectotype, Jamaica (10x).

264 JOHNSONIA, No. 37 Tricolia

D. and N. E.Schmidt); Big Pine Key; Summerland Key (both D. and N. E. Schmidt) ;
Middle Sambo Shoals, S of Boca Chica Key (T. McGinty); Boca Grande Key; Garden
Key (both FSM); Loggerhead Key (T. McGinty), both Dry Tortugas. Banama Is-
LANDS: Elbow Cay, Great Abaco Id. (R. Robertson); S of North Cay, in 2 fathoms;
off Nassau Harbour; Lyford Cay, all New Providence (all T. McGinty); Governors
Harbour, Eleuthera; Campdown, northern Cat Island (both MCZ). Cusa: Cabo San
Antonio; Cabo Cajon; off Los Arroyos; Dimas, in 4-5 fathoms; Santa Lucia, in 2-4
fathoms; La Esperanza; Cayo Arenas, in 2 fathoms; Cayo Levisa, in 2-3 fathoms;
Cabanas, all Pinar del Rio (all USNM): near Habana, Habana; Bahia de Cardenas,
Matanzas (both ANSP); off Cayo Fragoso; Caibarién, both Las Villas; off Punta Ale-
gre, Camagiiey; Guarda la Vaca, near Puerto Sama, Oriente (all MCZ). Jamaica:
(MCZ; AMNH: CNHM); Jack’s Bay, St. Mary (USNM); Port Antonio, Portland
(ANSP): Port Royal (USNM). Hispantota: Cap Haitien, Dépt. du Nord (MCZ);
Port Salut; Les Cayes; Aquin, all Dépt. du Sud; Saltrou, Dépt. de Ouest, all Haiti
(all USNM); Montecristi; El Canal, Cabo Macoris (both MCZ); Bahia de Samana
(USNM), all Dominican Republic. Purrro Rico: Aguadilla (G. Warmke); Arecibo;
Boca de Congrejos (both MCZ); Guanica (A MNH); Cabo Rojo (G. Warmke; MCZ):
Rineén (LSJU). Virern Istanps: St. Thomas (MCZ; USNM; CNHM); St. John
(MCZ). Lesser ANTILLES: Saint Martin (ANSP).

Tricolia affinis pterocladica, new subspecies
Plate 144, fig. 3; Plate 145, fig. 3

Description. Shell fairly solid, elongate, inflated and reaching 8 mm. in length. The
color pattern consists of wavy brown spiral lines which descend from the suture at an
angle of about 45°. There are usually white lines next to the brown lines. Below the
suture and at the periphery there are usually irregular, more or less axial, patches of
white. On the body whorl there are 7 or 8 of these. The ground color is light yellowish
brown. Occasionally the brown lines break up into dots on the lower part of the body
whorl. At the northern end of the range some specimens are red rather than brown.
Whorls 5, evenly rounded. Spire produced at an angle slightly less than 60°. Proto-
conch white, minute and depressed. Aperture elongate-ovate. Columellar callus thin.
Umbilicus reduced to a minute chink. Suture impressed. Post-nuclear whorls smooth
except for minute axial growth lines. Operculum tinged with brown at the margin.

length width
(large) 8.0 mm. 4.6mm. Miami Beach, Florida
4.6 3.1 Holotype, Boynton Beach, Florida
(juvenile) 9.5 1.9 Yamato Rocks, Florida

Types. ‘The holotype is in the Museum of Comparative Zoélogy, no. 215662. It was
collected by ‘Thomas L. McGinty on March 20, 1957, amongst Pterocladia growing on
rocks in shallow water at Boynton Beach, Florida. Paratypes are in the Museum of Com-
parative Zodlogy (no. 215663) and the United States National Museum, at Stanford
University, the Academy of Natural Sciences of Philadelphia (nos. 150755, 189846,
195923), the Chicago Natural History Museum (nos. 26488, 53118) and the private col-
lection of T. McGinty.

———— eee

Western Atlantic JOHNSONIA, No. 37 265

Remarks. This new subspecies can readily be distinguished from the nominate form
by the brown (rarely red) color, and, in particular, the color pattern of descending wavy
lines. From 7". tessellata, with which it has been confused, it differs in shape, the spire
being higher and the shell growing to a larger size. The spiral lines are far more irregu-
lar and descend more rapidly.

At Boynton Beach this subspecies lives amongst the red alga Pterocladia (P. ameri-
icana Taylor?) which grows on rocks in shallow water. I am indebted to Mr. Thomas L.
McGinty for collecting specimens of this alga and to Dr. William Randolph Taylor for
the identification. Inthe Florida Keys 7°. affinis affinis is known to live on Turtle Grass
(Thalassia). Thalassia does not live north of the Biscayne Bay area on the east coast of
Florida (Voss and Voss 1955). As the range of 7". affinis affinis is coextensive with that
of Thalassia in southern Florida it is here assumed that pterocladica is a geographically
segregated ecologic race which is adapted to live on Pterocladia rather than on Thalassia
and hence is able to live farther to the north than the nominate form. The range, how-
ever, of pterocladica does not coincide with the range of Pterocladia americana Taylor
(1943). This alga is found from North Carolina and Bermuda to Texas and the Carib-
bean. It is significant to observe in this connection that the ranges of 7". affinis affinis
and affinis pterocladica overlap slightly, the latter appearing sporadically in the northern
Florida Keys, and that there is barely any intergradation between the two. It is to be
hoped that field studies can be carried out to confirm or modify the suggestions made
here on the basis of the preliminary information available. Ticolia may possibly not be
as dependent on specific plant substrates as here implied.

Range. The southeast coast of Florida, from Fort Pierce to the Biscayne Bay area,
sporadically as far south as the northern Florida Keys (Grassy Key). ‘Three specimens
have also been collected off northwest Florida.

Specimens examined. Fioripa: Fort Pierce (D. and N. E. Schmidt; Mary Godwin);
Palm Beach (USNM; ML; G. and M. Kline); NE of Lake Worth, in 45 ft. (FSM):
Peanut Id., North Inlet, Lake Worth (T. McGinty); off Lantana, in 10 fathoms (FSM);
Boynton Beach (MCZ; T. McGinty; ANSP; CNHM); South Inlet, Lake Worth
(MCZ; FSM); Yamato Rocks (MCZ; FSM); Miami Beach (MCZ; ANSP; FSM;
USNM; U of M); Miami (MCZ; USNM); Fisher Id., Miami (T. McGinty); Norris
Cut (MCZ); Virginia Key (FSM); Bear Cut (D. Moore); Cape Florida, Biscayne
Key (FSM); Ragged Keys (USNM): Lower Matecumbe Key (MCZ); Grassy Key
(U of M); off Destin, near Fort Walton, in 14 fathoms (T.. McGinty).

Tricolia affinis beaui, new subspecies
Plate 144, fig. 4; Plate 145, fig. 5

Description. Shell solid, elongate, inflated and reaching 7.5 mm. in length. The color
pattern consists of irregular red spiral lines, each bordered by a white line, which descend
across the whorls from the suture at an angle of about 40°. The ground color is light
pink or pale orange. Below the suture and at the periphery there are red (occasionally
brown) patches; often these fuse to form irregular axial stripes. The red lines sometimes
break up into dots on the body whorl. Between the red patches below the suture there
are often white patches which are sometimes tinged with yellow. On the body whorl

JOHNSONIA, No. 37 Tricolia

266

there may be as many as 9 of these. Whorls 6, slightly flattened at the periphery. Spire
produced at an angle of 60°. Protoconch minute, white, depressed. Aperture oval. Colu-
mellar callus thick, white. Rimately perforate. Suture moderately impressed. Post-
nuclear whorls smooth except for minute spiral striae. Operculum fairly thin, white.

length width
7.0mm. 4.8mm. Holotype, Barbados, Lesser Antilles
3.2 2.4 Barbados, Lesser Antilles

Types. The holotype is in the Museum of Comparative Zodlogy, no. 215664. It was
collected by Mrs. F. G. Kellett at Bathsheba, Barbados. Paratypes are in the Museum
of Comparative Zoédlogy, no. 215665, and in the United States National Museum, the
Academy of Natural Sciences of Philadelphia, the American Museum of Natural His-
tory, the Chicago Natural History Museum, at Stanford University and the California
Academy of Sciences.

Remarks. This new subspecies is similar to the red phase of 7". affinis pterocladica
which appears only in the vicinity of Fort Pierce on the southeast coast of Florida. It
differs in reaching a larger size, and by having the subsutural patches smaller, more nu-
merous and often tinged with yellow. From the nominate form of the species it differs
in having the wavy red spiral lines which descend obliquely across the whorls.

Plate 144. Figs. 1-2. Tricolia thalassicola Robertson. Fig. 1. Holotype, near Elbow Cay, Great Abaco Id.,
Bahama Islands (11.5x). Fig. 2. Deep water form (juvenile), 30 fathoms, off Palm Beach, Florida (22x).
Fig. 3. Tricolia affinis pterocladica Robertson. Holotype, Boynton Beach, Florida (11x). Fig. 4. Tricolia
affinis beaui Robertson. Holotype, Bathsheba, Barbados, Lesser Antilles (10x). Fig. 5. Tricolia affinis cru-
enta Robertson. Holotype, Bahia do Flamengo, Ubatuba, Sao Paulo, Brasil (18x).

Western Atlantic JOHNSONIA, No. 37 267

The subspecific name is given in memory of Commandant Beau, an ardent collector
in Guadeloupe and Martinique over a century ago. It was through his efforts that many
of the more interesting marine mollusks of the Lesser Antilles were first discovered. For
his obituary see Journal de Conchyliologie 7, p. 393, 1858.

Range. The Lesser Antilles from Antigua to Tobago.

Specimens evamined. Lesser ANTILLES: Falmouth and English Harbour, Antigua
(USNM); Martinique (AMNH); Bathsheba, Barbados (MCZ); Barbados (USNM:
ANSP; LSJU); Pigeon Pt., Tobago (MCZ).

Tricolia affinis cruenta, new subspecies
Plate 144, fig. 5; Plate 145, figs. 4, 6

Tricolia (Tricolia) pygmaea ‘Philippi’ Haas 1953, Fieldiana: Zoology 34, pp. 204-205 (Ilha Grande, Rio de
Janeiro, Brasil); non Phasianella pygmaea Philippi 1848.

Description. Shell solid, oval, inflated and reaching 6.2 mm. in length. The color pat-
tern consists of rather large more or less square or oblong dark red spots, arranged in
regular spiral rows; occasionally some of the spots are white. The ground color is light
orange or white, tinged with red. Below the suture and at the periphery there may be
irregular white or red axial stripes. The spots are whitish on the white stripes and darker
red on the red stripes. Sometimes the subsutural and peripheral stripes fuse. Whorls 44,
evenly rounded and smooth. Spire usually produced at an angle of from 65° to 75°; apex
rounded. Protoconch minute, depressed and white. Aperture oval. Columellar callus
fairly thin, white. Umbilicus reduced to a chink. Suture impressed. Operculum dark
olive-green at the margin. Two specimens from Pernambuco, Brasil, are unusually elon-
gate, with the spire produced at an angle of about 50° (Plate 145, fig. 6). On these speci-
mens the red spots are less regular and have fused in patches here and there.

length width
(large) 6.2mm. 3.9mm. Pernambuco, Brasil
4.8 3.3 Puerto Cabello, Venezuela
4.0 3.0 Holotype, Ubatuba, Sao Paulo, Brasil
3.2 Doe Itajai, Santa Catarina, Brasil

Types. The holotype is in the Museum of Comparative Zoélogy, no. 215666. It was
collected at Bahia do Flamengo, Ubatuba, Sao Paulo, Brasil, by W. Narchi on January
7, 1956, and was donated by Joao de Paiva Carvalho. Paratypes are in the Museum of
Comparative Zodlogy, no. 215667, and in the Instituto Oceanografico, Sio Paulo, the
Academy of Natural Sciences of Philadelphia, and at Stanford University. Paratypes
from Sao Sebastiao and Villa Bella, Ilha de Sao Sebastiaio, both Sao Paulo, are in the
United States National Museum (nos. 150775, 180790, 364140).

Remarks. This new subspecies can be recognized by the regularly spaced large red
spots. The spire is usually lower than in other forms of 7°. affinis. It was incorrectly
identified by Haas (1953) as Phasianella pygmaea Philippi (1848). Philippi described
this from an unknown locality and figured it in 1858 (pl. 4, fig. 11). The spire is much
higher than is usual in crwenta, being produced at an angle of 55°, the suture is more

268 JOHNSONIA, No. 37 Tricolia

impressed and the aperture is circular rather than oval. Furthermore, the color pattern
is entirely different. The red spots are much smaller than in cruenta and, in addition,
pygmaea has large yellow flames below the suture and at the periphery. Pilsbry (1888,
p. 284) inadvertently synonymized Philippi’s name with 7” tessellata. The identity of
Philippi’s species still remains in doubt.

The Brasilian records of T'ricolia concinna C. B. Adams given by Lange de Morretes
(1949) and Gofferjé (1950) are no doubt based on this subspecies.

Specimens from Texas are probably Pleistocene or sub-Recent fossils. Some approach
T. affinis pterocladica in color pattern.

The subspecific name is a Latin adjective meaning stained or spotted with blood.

Range. Along the Caribbean coast of Central and northern South America, ranging
as far north as the Grenadines in the Lesser Antilles and south along the coast of Brasil
to the State of Santa Catarina. Sporadic on the shores of the western Gulf of Mexico.

Specimens examined. Texas: Rockport (J. Bequaert); Ransom Island, Aransas Bay
(D. Moore). Mexico: Veracruz (MCZ; ANSP); Dzilam de Bravo (ANSP); Yuca-
tan Channel (USNM). Honpuras: (USNM; ML). Panama: Bocas del Toro, Isla de
Col6n (T. McGinty; ANSP). Cotompia: Cartagena (CAS). Carippean IsLANDs:
Curacao; Bonaire (both ANSP). VENEzUELA: Puerto Cabello (MCZ). Lesser An-
TILLES: Admiralty Bay, Bequia, Grenadines (MCZ); Tobago (USNM); Buccoo Bay,
Tobago(ANSP). Brasiu: Fortaleza, Ceara (H. de Souza Lopes); Pernambuco (MCZ);
Ponta Verde, Maceié, Alagoas; Itapoan, Salvador, Bahia: Macaé (all H. de Souza
Lopes); Ilha Grande (CNHM), both Rio de Janeiro; Bahia do Flamengo, Ubatuba
(Joio de Paiva Carvalho); Sao Sebastiaéo; Villa Bella, [Ilha de Sao Sebastiéo; Guaruja,
near Santos (all USNM): Santos (H. de Souza Lopes; ANSP), all Sio Paulo; Cabe-
cudos, Itajai, Santa Catarina (H. de Souza Lopes).

Tricolia adamsi Philippi
Plate 143, figs. 3-4; Plate 145, fig. 2

Phasianella brevis C. B. Adams 1850, Contributions to Conchology, no. 4, p. 67 (Jamaica); non P. brevis
d’Orbigny 1842, etc.

Phasianella adamsi Philippi 1853, Systematisches Conchylien-Cabinet, Nirnberg, (2) 2, pt. 5, p. 27 [new
name for P. brevis C. B. Adams 1850, non P. brevis d’Orbigny 1842, ete. |.

Eulithidium minutissimum ‘C.B. Adams’ Dall 1908, Proceedings United States National Museum 34, p. 255

[nomen n udum| .

Description. Shell fairly thin, subglobose, inflated and reaching 3.8 mm. in length.
The color pattern consists of irregularly spaced, rather small, round or oval red dots on a
light pink ground color which is partially transparent, except below the suture where it
is pure white. ‘The margin of this white area is incised, so that there are lobes of white
extending down toward the periphery of each whorl. Between each lobe (of which there
are about 8 on the body whorl) the reddish white ground color is intensified. The white
subsutural area often becomes obscure on the body whorl. Interspersed between the red
dots are a few white dots, except at the periphery. There may also be patches below the
periphery where the ground color is darkened. Whorls 34 to 4, evenly rounded. Spire
low, produced at an angle of from 76° to 85°: apex rounded. Protoconch minute, de-

Western Atlantic JOHNSONIA, No. 37 269

Plate 145. Fig. 1. Tricolia affinis affinis C. B. Adams. Great Abaco Id., Bahama Islands. Fig. 2. Tricolia
adamsi Philippi. Guana Id., Virgin Islands. Fig. 3. Tricolia affinis pterocladica Robertson. Miami Beach,
Florida. Figs. 4,6. Tricolia affinis cruenta Robertson. Fig. 4. Puerto Cabello, Venezuela. Fig. 6. Elongate
form, Pernambuco, Brasil. Fig. 5. Tricolia affinis beaui Robertson. Barbados, Lesser Antilles (all 13x).

270 JOHNSONIA, No. 37 Tricolia

pressed, white or reddish purple. Aperture round to oval. Columellar callus obscure;
rimately perforate. Suture impressed. Post-nuclear whorls smooth except for fine spiral
striae on the penultimate whorls. Operculum oval, white, strongly convex exteriorly

and polished.

length width
(large) 3.8 mm. 3.0mm. Guana Id., Virgin Islands
3.7 2.8 Boca de Congrejos, Puerto Rico
2.8 2.4 Holotype of Phasianella brevis C. B. Adams
(juvenile) ee 1.3 Off Punta Alegre, Camagiiey, Cuba

Types. Turner (1956, pl. 21, fig. 4) selected a neotype of Phasianella brevis C. B.
Adams (Plate 148, fig. 4) from St. Thomas, Virgin Islands, believing that the types
were lost. This specimen is in the Museum of Comparative Zoédlogy, no. 186592. Sub-
sequently a single Jamaican specimen from the C. B. Adams collection has been re-
covered (Plate 143, fig. 3). Both specimens had been borrowed by Pilsbry from Amherst
College before the collection was obtained on exchange by the Museum of Comparative
Zoblogy in 1942. The neotype specimen was not returned until after the first paper on
C. B. Adams’ types was published (Clench and ‘Turner 1950) and the holotype was not
returned until after the supplement was published by Turner (1956). The holotype is now
in the Museum of Comparative Zodlogy, no. 215661. The neotype as a result is invalid.
The type locality is Jamaica.

Remarks. This species can be distinguished from 7’. affinis affinis, with which it com-
monly occurs and has been confused, by the smaller size, the lower spire, and the different
color pattern. The white subsutural area of adamsi is characteristic and the small red
spots are not consistently paired with white spots as they are in 7°. affinis affinis. The
present species varies little in color throughout its range. No information is available on
its ecology.

Dall (1889a) states that Phasianella brevis C. B. Adams “‘is merely a young specimen
of the shell he [C. B. Adams] had previously named T'urbo pulchella.”’ The specimens
seen by Dall must have been misidentified.

Three specimens in the United States National Museum are labeled ‘‘Hudlithidium
minutissima C. B. Ads.’’ Although Dall, as indicated in the synonymy above, has re-
ferred to this name, it was never published by C. B. Adams.

Range. Throughout the West Indies, from the Bahama Islands to the Lesser Antil-
les, and the coast of Central America (Honduras and Panama). Not present on the coast
of Florida.

Specimens evamined. BAHAMA IsLANDS: Elbow Cay, Great Abaco Id. (R. Robertson) ;
Mangrove Cay, Andros Id. (USNM). Off Nassau Harbour (T. McGinty); Treasure
Island (G.and M. Kline); Clifton Bluff (T. McGinty), all New Providence ; Campdown,
northern Cat Island (MCZ). Cusa: Dimas and Cabafias, Pinar del Rio (both USNM);:
near Habana (ANSP); Arenas de la Chorrera, both Habana; off Punta Alegre, Cama-
giiey; Guantainamo, Oriente (all MCZ). Jamaica: (MCZ; AMNH; CNHM); Jack’s
Bay, St. Mary; Port Royal (both USNM). Hispanioia: Les Trois Pavillons, Dépt.
du Nord-Ouest; Les Cayes; St. Louis du Sud; Aquin, all Dépt. du Sud; Saltrou,

Western Atlantic JOHNSONIA, No. 37 271

Dépt. de l'Ouest, all Haiti (all USNM); El Canal, Cabo Macoris (MCZ): Bahia de
Samana (USNM), both Dominican Republic. Puerro Rico: Aguadilla (G. Warmke);
Boca de Congrejos (MCZ). VirGin Is_tanps: St. Thomas (MCZ; USNM: U of M;:
CNHM); Guana Id., near Tortola (MCZ); St. Croix (G. Usticke). Lesser ANTILLES:
Saint Martin (ANSP); English Harbour, Antigua (USNM); Bathsheba, Barbados
(MCZ); Barbados (USNM; ANSP); Buccoo Bay, Tobago (ANSP). Honpvuras:
(USNM). Panama: Bocas del Toro, Isla de Colon (‘T. McGinty ; ANSP); Col6n(LSJU).

Tricolia thalassicola, new species
Plate 142, fig. 2; Plate 144, figs. 1-2; Plate 146

Phasianella umbilicata d’Orbigny 1842 [in] Ramon de la Sagra, Histoire de ]’Ie de Cuba, Mollusques 2,
p. 77, pl. 19, figs. 32-34 (Martinique, Guadeloupe, Jamaica and Cuba) [in part only]; non Littorina umbilicata
d’Orbigny 1840 [a Tricolia].

Description. Shell solid, oval, inflated and reaching 7.1 mm. in length. The color pat-
tern consists of a series of regularly spaced, large, light to dark brown, orange or olive-
green spots. There are about 7 pairs of axial to oblique flames of color on the body whorl
just below the suture and at the periphery; these are the same color as the spots but
darker. The spots on the flames are darkened. Between the flames the ground color is
cream-white; elsewhere it is cream. The spots are coalesced into oblique spiral bands in
some specimens (Plate 146, fig. 3). The spots are occasionally obscure and they then al-
ternate with pale patches. Whorls 6, evenly rounded. Spire usually produced at an angle
of from 65° to 77°; apex obtuse. Protoconch minute, depressed and white. Aperture
oval. Columella with a white callus. Umbilicus fairly narrow. Suture impressed. Post-
nuclear whorls with evenly spaced spiral striae; body whorl smooth except for minute
growth lines. Operculum white, strongly convex exteriorly and nearly smooth.

length width
(large) 7.1mm. 4.5mm. Cabo San Antonio, Cuba
4.3 3.3 Holotype, Great Abaco Id., Bahama Islands
(banded form) 2.9 2.3 Cabafias, Cuba
(deep water form) 2.4 1.8 Off Cape Lookout, North Carolina

Types. The holotype is in the Museum of Comparative Zoélogy, no. 218260. It was
collected by the author at Johnnie’s Cay, Drunken Cays, near Elbow [Little Guana]
Cay, Great Abaco Id., Bahama Islands, on July 28, 1953. Paratypes from the vicinity
of Elbow Cay are in the Museum of Comparative Zodlogy, no. 2138261, the United
States National Museum, the Academy of Natural Sciences of Philadelphia, the Amer-
ican Museum of Natural History, the Chicago Natural History Museum, at Stanford
University, the California Academy of Sciences, the Institute of Marine Biology, Puerto
Rico, and the private collection of Thomas L. McGinty.

Remarks. This species has been called Phasianella umbilicata d’Orbigny. However,
d’Orbigny’s figure and part of his description were based on 7°. affinis affinis. Untortu-
nately, the seven syntypes in the British Museum (Natural History) [no. 54,10. 4.283]
are thalassicola (one is figured here, Plate 142, fig. 2). The specimen figured originally by
d’Orbigny is evidently lost. It was deemed wiser to describe the present species as new

972 JOHNSONIA, No. 37 Tricolia

rather than merely to rename d’Orbigny’s homonym after selecting the syntype figured
here as the lectotype. These syntypes hence become invalid.

The species can readily be distinguished from 7’. affinis and 7. adamsi, with which it
has been confused, by the regularly spaced brown, orange, or olive-green spots, which
are relatively much larger than the red and the white spots of both these other species.
Also, 7. affinis is more elongate and grows to a larger size. The whorls of thalassicola are
more rounded, and the early whorls are evenly spirally striated, while in 7° affinis they
are smooth.

In arare color form of thalassicola the spots are coalesced so that the color pattern con-
sists of broad yellowish brown obliquely spiral bands on a cream ground color (Plate 146,
fig. 3). This form has been referred to as 7°. tessellata, but it is clearly distinguishable by
the shape of the shell, the spirally striated initial whorls and the thickness of the obliquely
spiral bands of color. Sometimes some of the spots near the base of the columella do not
coalesce, showing that this is merely a color form differing in no significant way from the
spotted form. It occurs sporadically in shallow water along the Florida Keys and in the
Caribbean.

A deep water form (Plate 144, fig. 2; Plate 146, fig. 2) occurs at both the northern
and southern ends of the range—North Carolina and Brasil—as well as off southern,
western, and northwestern Florida and off Yucatan, all in depths of from 10 to 35 fathoms.
Empty shells have been dredged as deep as 117 fathoms. This form was erroneously iden-
tified by Dall, in part, as Phasianella brevis d’Orbigny, an entirely different species, here
referred to the genus Gabrielona. The deep water form is much smaller than the shallow
water forms of thalassicola. The suture is more impressed and the whorls are more in-
flated. The characteristic spots of thalassicola are much smaller but are similarly arranged,
often with axial to oblique stripes below the suture and at the periphery. The initial
whorls are spirally striated as in the shallow water forms. This deep water form may pos-
sibly be a species distinct from thalassicola but its discontinuous and peripheral distribu-
tion suggests that this is not the case.

Plate 146. Tricolia thalassicola Robertson. Fig. 1. Great Abaco Id., Bahama Islands. Fig. 2. Deep water
form, off Cape Lookout, North Carolina. Fig. 3. Banded form, Cabafias, Pinar del Rio, Cuba (all 13x).

Western Atlantic JOHNSONIA, No. 37 273

This species normally lives on Turtle Grass (Thalassia testudinum Konig). The specific
name is derived from this fact. It may feed on the Thalassia itself, or, which is more
likely, onthe epiphytic algae. Little information has been published on the distribution
of Thalassia in the Western Atlantic except that by Ostenfeld (1927) and by Voss and
Voss (1955) on Florida. It does not live off North Carolina and it is unlikely that it
would grow in water deeper than 10 fathoms, so the deep water form of thalassicola
probably lives in another habitat. Thalassia does not live in Lake Worth, Florida, yet
specimens of thalassicola have been collected there, but they are small and do not show
the color pattern of spots clearly. The large specimens with prominent spots, so abundant
in the Bahama Islands and along the north coast of Cuba, live invariably on Thalassia.
They have been dredged in shallow water to a depth of 5 fathoms, being most abundant
where the sediment is calcareous. Variation in this species is probably due in part to
different plant foods.

Range. From off Cape Hatteras, North Carolina, throughout the West Indies, to off
Cabo de Sao Roque, Rio Grande do Norte, Brasil. There are no records from the west-
ern half of the Gulf of Mexico and the Atlantic coast of Central America and Colombia.
Thalassia, however, does live in these areas.

Specimens examined. NortH Carouina: Albatross, station 2596 (N. Lat. 35°08’30”;
W. Long. 75°10/00”) in 49 fathoms (dead); A/batross, station 2595 (N. Lat. 35°08/00” ;
W. Long. 75°05’30”) in 63 fathoms (dead), both off Cape Hatteras; Albatross, station
2617 (N. Lat. 33°37/30"; W. Long. 77°36’30’) in 14 fathoms (dead), off Cape Lookout
(all USNM). Fiorina: off Palm Beach in 30 fathoms (T. McGinty); Boynton Beach
(ANSP; T. McGinty); Hols, stations 49, 51, 70, 118, 124, in 10-85 fathoms, all off
Miami (all USNM); Fisher Id., Miami (T. McGinty); Hols, stations 8, 105, 107, 123,
142, 150, 306, in 22-100 fathoms (all dead), all off Fowey Light (all USNM); Turtle
Harbor (ANSP; U of M); Kolis, station 59, in 20 fathoms (dead), off Turtle Harbor
(USNM); off Carysfort Light, in 66-117 fathoms (dead); off The Elbow, Key Largo,
in 66-75 fathoms (dead; both MCZ); Upper Matecumbe Key (USNM); Conch Keys
(T. McGinty); off Sombrero Light, in 58 fathoms (dead; MCZ); Key West; Lolis,
station 44, in 50 fathoms (dead), off Key West (both USNM); Garden Key (FSM);
Loggerhead Key (USNM:;: FSM; T. McGinty), both Dry Tortugas; 90 miles SW of
Egmont Key, S of St. Petersburg, in 70 fathoms (dead); off Destin, near Fort Walton,
in 14 fathoms (dead; both T. McGinty). Bauama IsLanps: N of Little Abaco Id.
(MCZ): Drunken Cays, Elbow [Little Guana] Cay and Tilloo Cay, Great Abaco Id.
(all R. Robertson); Sweetings Village, Great Abaco Id. (MCZ); Great Isaac (U of M):
10 miles NE of North Point, North Bimini (R. Robertson); North Bimini and South
Bimini (both USNM); off Gun Cay (U of M); South Cat Cay (USNM; T. McGinty),
all Bimini Ids. ; South Riding Rock (R. Robertson); Mangrove Cay, Andros Id. (MCZ:;
USNM): Lyford Cay, S of North Cay in 2 fathoms and off Nassau Harbour (all T.
McGinty); North Cay (CNHM), all New Providence Id.; Galloway Landing, Long
Id.; Matthew Town, Great Inagua Id. (both MCZ). Cuna: Cabo San Antonio: Punta
del Tolete, in 2-8 fathoms; off Los Arroyos; Dimas, in 4—5 fathoms; Santa Lucia, in
2-4 fathoms; La Esperanza, in 2-8 fathoms; Cayo Arenas, in 2 fathoms: Cayo Levisa,
in 2-3 fathoms; Punta Colorado, in 2-3 fathoms; Cabanas, all Pinar del Rio (all USNM);

274 JOHNSONIA, No. 37 Tricolia

Arenas de la Chorrera, Habana; Bahia de Matanzas: Atlantis, station 2993 (N. Lat.
23°24’: W. Long. 80°44’) in 580 fathoms (dead), off Bahia de Cardenas, both Matan-
zas: off Cayo Fragoso, Las Villas; off Punta Alegre, Camagiiey; Guarda la Vaca, near
Puerto Sama, Oriente (all MCZ). Cayman IsLanps: Grand Cayman (MCZ; ANSP).
Jamaica: Montego Bay, St. James: Port Antonio, Portland (both USNM). Hispan-
ioLA: Bariadéle; Les Cayes: Lle-a-Vache, all Dépt. du Sud, Haiti (all USNM): Monte-
cristi(MCZ: ANSP): Bahia de Samana (USNM), both Dominican Republic. Purrro
Rico: Vieques (USNM). Virein Istanps: St. Thomas (USNM; ANSP); St. John
(MCZ). Lesser ANTILLES: Saint Martin (ANSP): Falmouth and English Harbour,
Antigua; Carlisle Bay, Barbados (all USNM): Admiralty Bay, Bequia, Grenadines
(MCZ). Carippean Istanps: Curacao (USNM: ANSP). Mexico: Dzilam de Bravo
(ANSP): off Cabo Catoche (USNM), both Yucatan. Brasit: Albatross, station 2758
(S. Lat. 6°59/30"; W. Long. 34°47'00”) in 20 fathoms, off Cabo de Sio Roque, Rio
Grande do Norte (USNM).

Tricolia bella MW. Smith
Plate 139, fig. 5; Plate 142, figs. 4-5; Plate 143, fig. 5; Plate 147, figs. 1-2

Turbo (2) pulchellus C. B. Adams 1845, Proceedings Boston Society of Natural History 2, p. 7 (Jamaica);
non Phasianella pulchella Récluz 1843 [a Tricolia.

Littorina adamsii Reeve 1857, Conchologia Iconica 10, Littorina, no. 85 (Jamaica) [*‘Littorina pulchella,
C. B. Adams, MS., Mus. Cuming”’ cited as a synonym]; non Phasianella adamsi Philippi 1853 [a Tricolia}.

Phasianella bella “Pilsbry’ M. Smith 1937, East Coast Marine Shells, p. 81, pl. 31, fig. 20 [mew name for
Tricolia pulchella C. B. Adams 1845, non Récluz 1843, in Pilsbry’s MS. This is listed as a synonym of bella

by Maxwell Smith].

Description. Shell solid, conical, more or less carinate, with spiral cords and reaching
5.2mm. in length. The color pattern consists of red, pink, brown or orange-yellow spots
or flames on a white ground color. Sometimes the shell is greenish or khaki-colored:
rarely it is unicolored. The flames usually form irregular axial stripes and the spots are
usually delimited by the width of the spiral cords. Whorls 44 to 5, angular in the mid-
dle, with the body whorl biangular, or rounded. Spire produced at an angle of from 59°
to 64°. Protoconch fairly large and prominent, flattened apically and either white or
bright purple. Aperture nearly round. Columellar callus obscure. Rimately perforate.
Suture impressed. The keel above the periphery is usually strongly developed while the
keel below on the body whorl is always less prominent: both are usually spotted with
white. Above and below these keels and between them there are fine irregularly beaded
spiral cords. In a smooth form of the species (Plate 147, fig. 2) the initial whorls are
threaded but on the evenly rounded body whorl the cords are obscure and the keels are
replaced by stripes on which the color pattern differs from that on the remainder of the
whorl. The upper keel can always be seen on the early post-nuclear whorls. The opercu-
lum is round-ovate, white, externally convex and threaded near the margin. The juvenile
operculum is perforate (Plate 139, fig. 5).

length width
(smooth form) 5.2mm. 3.3mm. Miami Beach, Florida
4.3 3.0 St. Thomas, Virgin Islands
3.0 2.3 Lectotype of Turbo (?) pulchellus C. B. Adams

(juvenile) 0.7 0.6 Pigeon Cays, Andros Id., Bahama Islands

Western Atlantic JOHNSONIA, No. 37 975

Types. The lectotype of Turbo (¢) pulchellus C. B. Adams (Plate 148, fig. 5), selected
by Clench and Turner (1950, pl. 40, fig. 9), is in the Museum of Comparative Zodélogy,
no. 156358. Lectoparatypes from Jamaica are in the Museum of Comparative Zoédlogy
(no. 186165), the Academy of Natural Sciences of Philadelphia (no. 6453) and the United
States National Museum (no. 64386). Two syntypes of Littorina adamsii Reeve (Plate
142, figs. 4-5) are in the British Museum (Natural History), nos. 1957.6.3.4-5. The
type locality is Jamaica.

Remarks. This species can readily be identified by its characteristic shape. It was iden-
tified by Pilsbry (1888, pl. 39, fig. 95), in part, as Phasianella umbilicata d’Orbigny, a
name here discussed in the Remarks on JT. thalassicola.

Maxwell Smith unintentionally renamed C. B. Adams’ homonym by using Pilsbry’s
manuscript name. The specimen he figures and describes is the smooth form from Florida.

The smooth form of this species (Plate 147, fig. 2) occurs with, and intergrades with,
the ribbed form and is most abundant from Miami to the Florida Keys and in parts of
the Bahama Islands, but also appears occasionally on the north coast of Cuba, on the
east coast of Guatemala and in the Lesser Antilles at Antigua. Even where it occurs
the smooth form is not uniformly distributed, for at many localities in the Florida Keys
and in the Bahama Islands only the ribbed form has been collected.

A specimen of the smooth form was collected by the author at Great Abaco Id., Ba-
hama Islands, on Thalassia in shallow water. These two variants are not ecologic forms,
however, because the ribbed form also lives on Thalassia, as well as in a variety of other
habitats, such as in calcareous gravel and beneath gorgonian holdfasts. The dimorphism
is probably entirely genetic. In general, the specimens mottled with green or khaki (both
smooth and ribbed forms) live on Thalassia, while those spotted with red are found
elsewhere.

Range. From Lake Worth south along the east coast of Florida, throughout the
Florida Keys and the West Indies, west to the coast of Central America and south at
least to the State of Alagoas, Brasil. Dall (1892) records the species from the Cedar
Keys on the west coast of Florida and as a Pliocene fossil in South Carolina.

Plate 147. Figs. 1-2. Tricolia bella M. Smith. Great Abaco Id., Bahama Islands. Fig. 1. Ridged form.
Fig. 2. Smooth form. Fig. 3. Tvricolia tessellata Potiez and Michaud. Barbados, Lesser Antilles (all 18x).

JOHNSONIA, No. 37 Tricolia

276

Specimens examined. FLoripa: North Inlet (T. McGinty); South Inlet (MCZ), both
Lake Worth; Bakers Haulover, Miami (ANSP); Miami Beach (ANSP; FSM); off
Government Cut, Miami; Holts, stations 49, 51, 70, in 10-80 fathoms (dead), off Miami
(all USNM); Fisher Id., Miami (‘T. McGinty); Ragged Keys; Elis, station 366, in
75-90 fathoms (dead), east of Ragged Keys; Molis, stations 8, 80, 306, in 25-100 fath-
oms (dead), off Fowey Light (all USNM); Sands Key, Biscayne Bay (MCZ; USNM);
Caesars Creek Bank, north of Key Largo (USNM); Key Largo (LSJU); Turtle Har-
bor, in 4-10 fathoms; Upper Matecumbe Key (both USNM); Teatable Key (T.
McGinty); Indian Key (USNM); Lower Matecumbe Key (USNM; CAS); Center
Key (LSJU); Conch Keys (T. McGinty; ANSP); Grassy Key (ANSP); Bonefish
Key (MCZ; ANSP; CNHM; LSJU; U of M; FSM); Littl Duck Key; Misseur
Key (both MCZ); Bahia Honda Key (USNM; ANSP); Noname Key (LSJU); Big
Pine Key (MCZ; ANSP); Newfound Harbor Key (USNM); Torch Key (ANSP);
Summerland Key (D.and N. E.Schmidt); Loggerhead Key (USNM); Sugarloaf Key;
Boca Chica Key (both ANSP); Middle Sambo Shoals, S of Boca Chica Key
(T. McGinty); Key West (MCZ; USNM; ANSP; LSJU; U of M); Boca Grande
Key (USNM). Garden Key (USNM; FSM); Loggerhead Key (USNM; T. McGinty);
Sand Key; Bush Key (both FSM), all Dry Tortugas. Banama Isitanps: N of Little
Abaco Id.; Whale Cay Channel; Sweetings Village, both Great Abaco Id. (all MCZ);
Elbow [Little Guana] Cay; between Channel Cay and Wilson City, both Great Abaco
Id. (both R. Robertson); Great Isaac (U of M). 5 miles NE of North Point, North Bi-
mini (R. Robertson); North Bimini (USNM); South Bimini (MCZ; USNM); South
Cat Cay (USNM; T. McGinty), all Bimini Ids.; South Riding Rock; Deep Creek,
Andros Id. (both R. Robertson); Pigeon Cays (T. McGinty) and Mangrove Cay (MCZ;
USNM), Andros Id. ; Clifton Pt., Lyford Cay, S of North Cay, and off Nassau Harbour,
all New Providence Id. (all T. McGinty); Simms; Galloway Landing, both Long Id.
(both MCZ); Acklins Id.(USNM). Cuzsa: off Los Arroyos; Dimas; Cayo Arenas; Cayo
Levisa; Bahia Honda; Cabanas, all Pinar del Rio (all USNM); Arenas de la Chorrera,
Habana (MCZ; CNHM); Bahia de Matanzas; Varadero (both MCZ); Bahia de Car-
denas (USNM), all Matanzas; off Cayo Fragoso, Las Villas; off Punta Alegre, Cama-
giiey; Guarda la Vaca, near Puerto Sama; Guantanamo, both Oriente (all MCZ).
Jamaica: Montego Bay, St. James; Jack’s Bay, St. Mary (both USNM); Port Royal
(MCZ; USNM): Mouth of Rio Cobre, St. Catherine(USNM). Hispanioia: Les Trois
Pavillons, Dépt. du Nord-Ouest; Bariadéle; Coteaux; Les Cayes; Aquin, all Dépt. du
Sud; Saltrou, Dépt. de l’Ouest, all Haiti (all USNM); Montecristi; Puerto Plata; El
Canal, Cabo Macoris (all MCZ); Bahia de Samana (USNM), all Dominican Republic.
Purerro Rico: Rineén (LSJU); Aguadilla; Boca de Congrejos (both G. Warmke).
VirGiIn Istanps: St. Thomas (USNM; ANSP); St. Croix (G. Usticke). LrssEr AN-
TILLES: Falmouth and English Harbour, Antigua (USNM); Martinique (AMNH);
Crane Pt., Barbados (USNM); Barbados(MCZ; ANSP; LSJU); Pigeon Pt. (MCZ);
Buccoo Bay (MCZ; ANSP), both Tobago. GuarEmaALa: Livingston and Puerto Bar-
rios (ANSP). Honpuras: Isla Utila (USNM). Panama: Bocas del Toro, Isla de
Colon; Pina, west of Colén; Col6n (all T. McGinty). Brasti: Maceié, Alagoas (H. de
Souza Lopes).

Western Atlantic JOHNSONIA, No. 37 277

Tricolia tessellata Potiez and Michaud
Plate 142, fig. 3; Plate 143, fig. 6; Plate 147, fig. 3

Phasianella tessellata ‘Beck’ Potiez and Michaud 1838, Galerie des Mollusques, Paris, 1, p. 312, pl. 29, figs.
7-8 (Martinique).

Phasianella minuta Anton 1839, Verzeichniss Conchylien, Halle, p. 60; non P. minuta J. Sowerby 1817 [is
a synonym of P. tessellata Potiez and Michaud according to Philippi 1853, p. 19].

Phasianella zebrina d’Orbigny 1842[in] Ramon de la Sagra, Histoire de l’Ile de Cuba, Mollusques 2, p. 78,
pl. 19, figs. 35-37 (Guadeloupe).

Phasianella tesselata C.B. Adams 1850, Contributions to Conchology, no. 4, pp. 67-68 (Jamaica) [misspelled
tessellata in republication by Clench and Turner 1950, p. 351).

Phasianella lipeata ‘Dkr.’ Krebs 1864, The West-Indian Marine Shells, p. 79 (republished by Clench,
Aguayo and Turner 1947, Revista de la Sociedad Malacolégica 5, p. 109) [momen nudum; cited as a synonym
of P. tesselata C. B. Adams ‘‘according to specimens in the collection of A. H. Riise’’].

Description. Shell solid, smooth, oval, inflated and reaching 5.2 mm. in length. The
color pattern consists of regularly spaced spiral lines which descend obliquely, arising
from the suture at an angle of about 15°. There are irregular axial streaks between these,
often in subsutural and peripheral patches. The lines and streaks are red or brown (some-
times with an olive-green tint) on a pale cream or pink ground color with irregular white
spots adjacent to them. The entire shell is sometimes brownish black, with the oblique
spiral lines darkest. Whorls 5, convex, slightly flattened or even concave above the pe-
riphery. Apex obtuse and the spire produced at an angle of from 72° to 76°. Protoconch
minute, red or purple. Aperture oval. Columella with a white callus. Umbilicus nar-
row. Suture slightly impressed. Body whorl often spirally striated, the striae deepest
just below the suture. Operculum white, tinged marginally with dark green and brown,
spirally striated internally; radially striated externally, most strongly near the margin.

length width
5.2mm. 4.5mm. Jamaica
3.7 3.0 Barbados
(juvenile) 1.5 1.4 Aquin, Haiti

Types. According to Gaillard (in Kaas, Basteria 21, p. 84, 1957) the Potiez and Mi-
chaud types formerly in the Musée de Douai, Dépt. du Nord, France, have disappeared.
Hence there are now no type specimens of Phasianella tessellata Potiez and Michaud. The
whereabouts of Anton’s types is unknown to the author. Seven syntypes of Phasianella
zebrina ad’ Orbigny (one is illustrated here, Plate 142, fig. 3) are in the British Museum
(Natural History), no. 54.10.4.284. The lectotype of Phasianella tesselata C. B. Adams
(Plate 143, fig. 6), selected by Clench and Turner (1950, pl. 36, fig. 12), is in the Mu-
seum of Comparative Zodlogy, no. 186067. Lectoparatypes from Jamaica are in the
Museum of Comparative Zoélogy (nos. 119808, 186068), the Academy of Natural Sci-
ences of Philadelphia (no. 6463) and in the United States National Museum (nos. 6418,
54762, 64387). The type locality of P. tessellata Potiez and Michaud is Martinique,
Lesser Antilles.

Remarks. This abundant species is easily recognized by the oblique spiral lines and the
shape of the shell. It is highly variable in coloration; some specimens are red, others are
light brown and a few are brownish black. The light brown form is abundant in Jamaica

978 JOHNSONIA, No. 37 Tricolia

but decreases in frequency to the east and south. In the Lesser Antilles where the red
form is abundant the brown form is searce. The red form is infrequent in Jamaica. The
brownish black form appears sporadically throughout the range. Specimens from Vene-
zuela, Colombia and Panama are of all three color forms and are strongly pigmented.

The differences between the present species and the banded form of 7°. thalassicola are
discussed in the Remarks under the latter species.

Records of this species in Florida are based on 7°. affinis pterocladica.’ No specimens
have been seen by the author from the Bahama Islands,’ Cuba or the northern portion
of Haiti. It probably does not live in these areas. The Brasilian record of T'ricolia minuta
Anton given by Lange de Morretes (1949), which presumably is based on this species,

is questionable.

Range. Jamaica, Hispaniola, Puerto Rico, south through the Lesser Antilles to Trin-
idad and west along the coast of northern South America to Panama.

Specimens examined. JAMAICA: Montego Bay, St. James; Jack’s Bay, St. Mary; Port
Morant, St. Thomas (all USNM); Port Royal (MCZ; USNM):; Mouth of Rio Cobre;
Old Harbour, both St. Catherine (both USNM). Hispanioia: Port Salut; Torbeck;
Les Cayes: St. Louis du Sud; Aquin, all Dépt. du Sud; Saltrou, Dépt. de l'Ouest, all
Haiti (all USNM). Montecristi(MCZ; ANSP); Puerto Plata; El Canal, Cabo Macoris
(both MCZ): Bahia de Samand (USN M), all Dominican Republic. PuErro Rico: Agua-
dilla (G. Warmke); Arecibo; Boca de Congrejos (both MCZ); Guaniquilla (LSJU);
Mayagiiez (USNM); off Mayagiiez, in 40-100 ft.; Rincon (both G. Warmke); Rincén
Lighthouse; Bahia de Afiasco; Pta. Arenas, Vieques (all MCZ). Virein Istanps: St.
Thomas (USNM: AMNH): Guana Id., near Tortola (MCZ); St. Croix (G. Usticke).
Lesser ANTILLES: Saint Martin (ANSP): Falmouth, Antigua (USNM); Guadeloupe
(MCZ: ANSP): Martinique (AMNH): Bathsheba, Barbados (MCZ: USNM): Bar-
bados (ANSP; AMNH; LSJU; CNHM): Anse Fourmi, Tobago; Carenage, Trini-
dad (both MCZ). VENEzuELA: Maiquetia, Federal District (ANSP). CoLomepia: Car-
tagena (CAS). Panama: Bocas del Toro, Isla de Col6én (T. McGinty).

‘M. Smith (1937) identified both species as tessellata. One of the specimens he illustrates is 7. affinis ptero-
cladica (pl. 29, fig. 18a) and the other is true fessellata (pl. 29, fig. 18b). The latter could not have been col-
lected in Florida.

* Ford’s record (1944) is erroneous.

Western Atlantic JOHNSONIA, No. 37 279

Notes

The synonymies are given here of two Eastern Pacific species which are the types of
Eulithidium Pilsbry and Usatricolia Habe, here considered synonyms of T'ricolia, s.s.

Tricolia variegata Carpenter
Plate 138, fig. 6; Plate 148, fig. 1

Eucosmia variegata Carpenter 1864, Annals and Magazine of Natural History (3) 13, p. 475 (Cape San
Lucas [Baja California, Mexico] ).

Eulithidium typicum Dall 1908, Proceedings United States National Museum 34, p. 255 [new name for Eu-
cosmia variegata Carpenter 1864, non Phasianella variegata Lamarck 1822}.

Phasianella (Eulithidium) typica Dall. Strong 1928, Proceedings California Academy of Sciences (4) 27
p. 194, pl. 10, figs. 12-13.

This species is the type of Hulithidium Pilsbry. It is to be observed that the spire is
not particularly low. It is known only from Cape San Lucas and Bahia Magdalena, Baja
California, Mexico. The holotype is in the United States National Museum, no. 11836.

Tricolia compta Gould
Plate 138, fig. 4; Plate 148, fig. 2

Phasianella compta Gould 1855, Appendix to the Preliminary Geological Report of William P. Blake (Pal-
aeontology). Explorations and Surveys for a Railroad Route from the Mississippi River to the Pacific Ocean
(War Department), U.S. House of Representatives Doc. 129, 33d. Congress, Ist. Session, pp. 22, 25 (San
Diego, California) [published as a separate but not included in the Congressional Reports]; ‘Gould, ms.’
Carpenter 1856 (May), Catalogue of the Reigen Collection of Mazatlan Mollusca in the British Museum, pp.
225-226 (San Diego and Santa Barbara, California; Mazatlan, Mexico); ‘Gould, MS.’ Carpenter 1857 (Jan-
uary 7), Proceedings Zoological Society of London 24, p. 204 (Santa Barbara and San Diego); Gould 1857,
Pacific Railroad Reports 5, pt. 2, pp. 330, 333-334, pl. 11, figs. 25-26 (San Diego).

Plate 148. Fig. 1. Tricolia variegata Carpenter. Bahia Magdalena, Baja California, Mexico (21x). The type
species of Eulithidium Pilsbry. Specimen collected and identified by A. M. Strong (CAS Loe. 24036). Fig. 2.
Tricolia compta Gould. Ideotype, Santa Barbara, California (11x). The type species of Usatricolia Habe.

280 JOHNSONIA, No. 37 Notes

Phasianella (Tricolia) compta Gould var. producta Dall 1908, Proceedings United States National Museum

34, p. 256 [refers to Pilsbry 1888, pl. 39, fig. 69].
Phasianella (Tricola) compta Gould. Strong 1928, Proceedings California Academy of Sciences (4) 17, p.

191, pl. 10, fig. 1.

This species is the type of Usatricolia Habe. It is apparently restricted to the coast of
California and northern Baja California. The specimen figured is an ideotype mentioned
by Carpenter from Santa Barbara. The location of Gould’s holotype of this species is
unknown. According to Palmer (1951) two specimens from Mazatlan (‘*Tablet 1086”
listed by Carpenter in 1856) are in the New York State Museum. These are probably a

different species.

Phasianella bicarinata Poulsen

Phasianella bicarinata ‘d’Orbigny’ Poulsen 1878, Catalogue of West-India Shells, Copenhagen, p. 13 [nomen

nudum}; non P. bicarinata Dunker 1846.

This was probably a misidentification of T7icolia bella. T. bicarinata Dunker is a sim-
ilar South African species. ;

Phasianella concolor C. B. Adams
Phasianella concolor C. B. Adams 1850, Contributions to Conchology, no. 4, p. 68 (Jamaica).

This is a synonym of Assiminea auberiana (d’Orbigny), as was first shown by Dall
(1892). The lectotype, selected by Clench and Turner (1950, pl. 36, fig. 3), is in the
Museum of Comparative Zodlogy, no. 186022; so also are lectoparatypes, no. 186023.
Pilsbry (1888, pp. 171, 275, pl. 45, fig. 1) erroneously applied this name to the dark
brown form of T'ricolia tessellata.

Phasianella tessellata Anton

Phasianella tessellata Anton 1839, Verzeichniss Conchylien, Halle, p. 61 (South America); non P. tessellata
Potiez and Michaud 1838.

As suggested by Philippi (18538, p. 82) this is probably a Littorina.

Dall (1889b) records both *‘Phasianella umbilicata Orbigny’* and ‘*P. pulchella C. B.
Adams”* [= Tricolia bella M. Smith] from Bermuda. Two species of T'ricolia are also
listed from the island by Tristram (1862). I have seen no authentic Bermuda specimens.

Western Atlantic JOHNSONIA, No. 37 281

REFERENCES

Adams, H. and A. 1854, The Genera of Recent Mollusca, London, 1, pp. 389-390.
Aguayo, C. G. 1945, Revista de la Sociedad Malacologica 3, p. 84.

Béggild, O. B. 1930, Det Kongelige Danske Videnskabernes Selskabs Skrifter, Naturvidenskabelig og Math-
ematisk Afd. (9) 2, p. 301, pl. 9, fig. 5.

Buequoy, E., P. Dautzenberg and G. Dollfus 1884, Mollusques Marins du Rousillon, Paris, 1, pp. 337-339.
Clark, W. 1855, British Marine Testaceous Mollusca, London, pp. 320-322, 516-517.

Clench, W. J. and R. D. Turner 1950, Occasional Papers On Mollusks, Harvard Univ., 1, no. 15.
Cossmann, M. 1889, Annales de la Société Royale Malacologique de Belgique 24, pp. 3-4.

Cossmann, M. 1918, Essais de Paléoconchologie Comparée, Paris, livraison 11, pp. 156-167.

Cotton, B. C. and F. K. Godfrey 1938, Malacological Society of South Australia Publication No. 1, p. 9.
Cotton, B.C. 1945, Transactions Royal Society of South Australia 69, pp. 163-165.

Crespin, I. 1926, Proceedings Royal Society of Victoria (2) 38, p. 119, pl. 9, figs. 16-17.

Dall, W. H. 1889a, Bulletin of the Museum of Comparative Zodlogy 18, p. 351, pl. 19, fig. 10b.

Dall, W. H. 1889b, United States National Museum Bulletin 37, pp. 158-159, pl. 19, fig. 10b.

Dall, W. H. 1892, Transactions Wagner Free Institute of Science of Philadelphia 3, pt. 2, pp. 347, 381.
Dall, W. H. 1908, Proceedings United States National Museum 34, p. 255.

Dall, W. H. 1915, United States National Museum Bulletin 90, pp. 94-95, pl. 12, fig. 5.

Davies, A. M. 1934, Tertiary Faunas, London, 2, pp. 104-105.

Eberhard, E. 1865, Uber die Schneckenzungen. Programm der Herzogleich realschule zu Coburg, p. 16,

pl. 5, fig. 98. Not seen.
Finlay, H. J. 1926, Transactions and Proceedings of the New Zealand Institute 57, p. 368.
Forbes, E. 1844, British Association for the Advancement of Science Report of the Thirteenth Meeting, p.138.

Forbes, E. and S. Hanley 1849-50, History of British Mollusca, London, 1, pl. DD, figs. 5, 5a; 2, pp.
537-540.

Ford, P. D. 1944, A Complete List of Bahamian Shells, p. 9. Privately printed: Nassau, Bahama Islands.

Fretter, V. 1955, Proceedings Malacological Society of London 31, pp. 159-162.

Gabb, W. M. 1873, Transactions American Philosophical Society (n.s.) 15, p. 240.

Gardner, J. 1947, United States Geological Survey Professional Paper 142-H, pp. 608-613, pl. 40, figs.
6-8, 13.

Gatliff, J. H. and C. J. Gabriel 1908, Proceedings Royal Society of Victoria (n.s.) 21, p. 366, pl. 21, figs.
O-10:

Gofferjé, C. N. 1950, Arquivos do Museu Paranaense, Curitiba, Brasil, 8, p. 232.

Gray, J. E. 1847, Proceedings Zoological Society of London 15, p. 144.

JOHNSONIA, No. 37 References

28%

i)

Gray, J. E. 1854, List of the shells of Cuba in the collection of the British Museum, pp. [iii], 22-23.

Guppy, R. J. L. and W. H. Dall 1896, Proceedings United States National Museum 19, pp. 322-323.
Haas, F. 1953, Fieldiana: Zoology 34, pp. 204-205.

Habe, T. 1956, Venus 19, pp. 95-96, figs. 1-2.

Harris, G. F. 1897, Catalogue of Tertiary Mollusca in the Department of Geology British Museum (Natural
History). Part I. The Australian Tertiary Mollusca, p. 275.

Ino, T. 1955, American Malacological Union Annual Reports for 1955, p. 30.

Iredale, T, 1924, Proceedings of the Linnean Society of New South Wales 49, pp. 230-232.

Jeffreys, J. G. 1865, British Conchology, London, 3, pp. 337-341, pl. 8, fig. 1.

Kiener, L.-C. 1847, Spécies des Coquilles Vivantes, Paris, 10, Genre Phasianelle, pp. 1-11, pls. 1-5.
Kuroda, T. and T. Habe 1954, Venus 18, pp. 86, 93-94, figs. 2, 5, 6.

Lange de Morretes, F. 1949, Arquivos do Museu Paranaense, Curitiba, Brasil, 7, p. 62.

Lebour, M. V. 1937, Journal Marine Biological Association of the United Kingdom 22, pp. 109, 110, 124,

fig. 1m.

Mansfield, W. C. 1937, State of Florida Department of Conservation Geological Bulletin 15, p. 180.

Martin, K. 1916, Sammlungen des Geologischen Reichs-Museums in Leiden (2) 2, pt. 6, p. 260, pl. 3, figs.

85, 85a.
Olsson, A. A. 1922, Bulletins of American Paleontology 9, pp. 332-333.
Ostenfeld, C. H. 1927, Die Pflanzenareale 1, pt. 3, pp. 35-36, map 22.
Palmer, K. V. W. 1947, Memorias de la Sociedad Cubana de Historia Natural 19, p. 96.
Palmer, K. V. W. 1951, New York State Museum Bulletin 342, p. 35.
Pelseneer, P. 1888, Bulletin Scientifique de la France et de la Belgique 19, p. 190.

Pelseneer, P. 1891, Sur l’oeil de quelques Mollusques Gastropodes. Annales Société Belge de Microscopie
16, p. 66. Not seen.

Pelseneer, P. 1899, Mémoires Couronnés par ]’Académie Royale des Sciences de Belgique 57, pt. 3, p. 46.
Philippi, R. A. 1848, Zeitschrift fir Malakozoologie 5, p. 18.

Philippi, R. A. 1853, Systematisches Conchylien-Cabinet, Niirnberg, (2) 2, pt. 5, pp. 1-32, pls. 1-5.
Pilsbry, H. A. 1888, Manual of Conchology, Philadelphia, (1) 10, pp. 162-184, pls. 37-40, 45, 60-61, 64.
Pilsbry, H. A. 1917, Proceedings Academy of Natural Sciences of Philadelphia 69, p. 207.

Priolo, O. 1953, Atti della Accademia Gioenia di Scienze Naturali in Catania (6) 8, pp. 109-113.

Quoy, J. R. C. and J. P. Gaimard 1832-35, Voyage de 1’Astrolabe, Zoologie, Atlas, pl. 59, figs. 1-2; text,
1834, 3, pp. 233-238.

Reeve, L. A. 1862, Conchologia Iconica 13, Phasianella, pls. 1-6.

Risbec, J. 1940, Annales des Sciences Naturelles, Zoologie, (11) 2, pp. 282-286, pl. 6, figs. 62-64, 66-70,

ple 7, Mes. Ta, 17.

>

Western Atlantic JOHNSONIA, No. 37 283

Robertson, R. 1957, Journal Washington Academy of Sciences 47, pp. 316-319, figs. 1-3.
Roissy, F. de 1805, Histoire Naturelle Générale et Particuliére des Mollusques, Paris, 5, p. 330.
Smith, M. 1937, East Coast Marine Shells, Ann Arbor, Mich., p. 81, pls. 29, 31.

Sowerby, G. B. II 1884, Thesaurus Conchyliorum 5, pp. 149-152, pls. 475-476.

Strong, A. M. 1928, Proceedings California Academy of Sciences (4) 17, pp. 187-202, pl. 10.

Taylor, W. R. 1943, Papers Michigan Academy of Sciences (Botany and Forestry) 28, pp. 154-156, pl. 4,
fig. 1.

Thiele, J. 1924, Mitteilungen Zoologischen Museum in Berlin 11, pp. 63-64, 72, fig. 23.

Thiele, J. 1929, Handbuch der systematischen Weichtierkunde, Jena, 1, pp. 70-71, fig. 49 (see correction
published in 1935, 2, p. 1154).

Torr, C. M. 1914, Transactions and Proceedings Royal Society of South Australia 38, p. 364, pl. 19, figs.

5a, ob.
Tristram, H. B. 1862, Proceedings Zoological Society of London for 1861, p. 405.
Troschel, F. H. 1878, Gebiss der Schnecken, Berlin, 2, pp. 200-203, pl. 18, figs. 9-16.
Turner, R. D. 1956, Occasional Papers On Mollusks, Harvard Univ., 2, no. 20, p. 136, pl. 21, fig. 4.
Voss, G. L. and N. A. Voss 1955, Bulletin of Marine Science of the Gulf and Caribbean 5, pp. 220-221.
Wenz, W. 1938, Handbuch der Paliozoologie, Berlin, 6, pt. 2, pp. 361-364, figs. 853-862.
Woodring, W. P. 1928, Carnegie Institution of Washington Publication 385, pp. 418-421, pl. 34, figs. 7-11.

Woodring, W. P. 1957, United States Geological Survey Professional Paper 306-A, pp. 65-66, pl. 15, figs.
1-2, pl. 17, fig. 47.

* *K *k

The Museum Boltenianum or the Bolten Catalogue

Probably no one book and particularly one so small has brought forth so much com-
ment or caused so much controversy in the field of malacology as the 199 page catalogue
of the Museum Boltenianum which was published in 1798. The Bolten Catalogue, as it
is commonly called, was originally a manuscript classification which Dr. J. F. Bolten of
Hamburg, Germany, had worked out for his large shell collection. It is believed that he
had intended to publish his system of classification but he died before it was completed.
Later, P. F. Réding, a book dealer and student of malacology, added to the manuscript
references to Gmelin 1791, the Conchylien-Cabinet and other publications in order that
the names might be identified. This was done at the request of the family and the cata-
logue was published and offered for sale, partly as a tribute to the late Doctor but also
with the hope that the collection would be purchased as a unit by some museum or
scholar. Dr. Bolten had disagreed with much that had been done in malacology up to
that time and so, diverging greatly from the Linnean system, instituted a multitude of

284 JOHNSONIA, No. 37 Museum Boltenianum

new generic and specific names. Thus the catalogue consists simply of a list of genera
and species with references to other publications.

Though offered for sale, the catalogue apparently was not widely distributed and was
little used, and Lamarck’s classification and names which were published in 1799 were
followed for many years. Of the early workers who referred to the Museum Boltenianum,
QO. A. Morch and H. and A. Adams were the most important. It was in 1906 that
Dall brought the Museum Boltenianum to the fore and in the same year Sherborn and
Sykes brought out a photographic facsimile of the catalogue, thus making it more avail-
able. This was followed by a lively discussion as to whether or not the catalogue was
truly a publication or only a sales catalogue, and an equally ardent controversy waged
concerning the author—was it Bolten or Réding? It is generally considered today that
since it was Réding who added the necessary references to identify Bolten’s names he
should be considered the author.’ When in 1926 the International Commission on Zoo-
logical Nomenclature made a ruling (Opinion 96) that the Museum Boltenianum was
an acceptable publication in systematic zoology, the names instituted in it had priority
over those of Lamarck and several other workers. Consequently many old and well-known
names were put into synonymy and most malacologists, following the ruling of the Com-
mission, set themselves to the task of revising various groups, using Réding’s names.
Now, after more than 30 years of recognition, the names instituted in the Museum
Boltenianum have come into general use.

The following partial bibliography is given for those who wish to go further into the
history and discussion concerning this famous little book.

Dall, W.H. 1906, Early History of the Generic Name Fusus. Journal of Conchology 11, pp. 289-297.

Dall, W.H. 1915, An Index to the Museum Boltenianum. Smithsonian Institution Publication, no. 2360,
pp. 1-64.

International Commission on Zoological Nomenclature 1926, Opinion 96. Smithsonian Miscellaneous Collec-

tions 73, no. 4, pp. 16-18.
Iredale, T. 1939, Mollusca Part I. Great Barrier Reef Expedition Scientific Reports 5, no. 6, pp. 373-374.
Rehder, H.A. 1945, A Note on the Bolten Catalogue. Nautilus 59, pp. 50-52.

Réding, P.F. 1798, Museum Boltenianum, Hamburg, pp. viiit+1—199 [photographic facsimile by Sherborn
and Sykes, London 1906].

Réding, P.F. 1819, Museum Boltenianum, Second Edition, pp. 1-156, 4 plates. [This edition was prepared
specifically for the sale of the Bolten collection at auction. |—RUTH D. TURNER

' Article 21, International Rules of Zoological Nomenclature (1905).

JOHNSONIA :

— Published by

Tue DEPARTMENT OF MOLLUSKS niece

Museum of Comparative Zoédlogy, Harvard University
Cambridge, Massachusetts

JUNE 28, 1958 PINNIDAE VOL. 3, NO. 38

THE FAMILY PINNIDAE IN THE WESTERN ATLANTIC
BY
Rutru D. TurNerR anp JoseErpH ROSEWATER

The Pinnidae is a small family of highly specialized bivalves which belongs to the order
Anisomyaria, that is, bivalves which have a large posterior adductor muscle, while the
anterior adductor is small or lacking. They are world-wide in distribution in warm-
temperate and tropical seas. Most species are found in sandy and muddy areas, from the
low water line to depths of about 10 to 25 fathoms. They are wedge- or fan-shaped _ bi-
valves which are equivalve but inequilateral. The umbos are terminal and are located at
the very narrow, anterior end which is buried in the substratum. Young specimens, at the
time of settling, are normal, equilateral bivalves as is shown in Plate 150, figs. 1-6. How-
ever, as growth is almost entirely in a posterior direction, the shell very quickly becomes
triangular in shape and the umbos become terminal. As the animal begins its burrowing

Plate 149. Cross section of the prismatic layer of Pinna carnea Gmelin showing the irregular prisms with con-
chiolin surrounding them (470x).
285

286 JOHNSONIA, No. 38 Pinnidae

activities the anterior end of the shell, being downward, becomes eroded. Consequently,
specimens with the embryonic valves still attached are extremely rare. As the shell grows,
the body of the animal as well as both the anterior and posterior adductor muscles must
move posteriorly (i.e., up). The anterior end of the shell is constantly worn away and the
mantle deposits septa across the shell to protect the anterior adductor muscle (Plate 154,
figs. 2, 4). Specimens are often found with a series of such septa in the narrow ends of
the valves in front of the small anterior adductor muscle scar.

SHELL STRUCTURE

The shell is very simple in structure and is composed generally of only two layers, the
periostracum being extremely thin or entirely absent. The outer prismatic layer is cal-
citic and composed of very large crystals produced in a framework of conchiolin and set
at right angles to the surface of the valves. These prismatic crystals, the largest pro-
duced anywhere in the Recent Mollusca, may be readily seen with the aid of a hand lens

Plate 150. Embryonic valves of Pinnidae. Figs. 1-2. Pinna carnea Gmelin from reef northeast of North Point,
Elbow Cay, Great Abaco, Bahama Islands. Figs. 3-4. Pinna rudis Linné from the Mediterranean. Figs.
5-6. Atrina (Servatrina) seminuda Lamarck from off Tobago, Lesser Antilles in 36 fathoms.

Western Atlantic JOHNSONIA, No. 38 287

and in older specimens are often sufficiently large to be seen without magnification.
Transverse and longitudinal -sections of the prismatic layer of Pinna carnea are shown in
Plate 149 and Plate 151, figs. 1-2. That portion of the shell posterior to the large pos-
terior adductor muscle is composed only of this prismatic layer and, because of its high
organic content, is slightly flexible in living specimens. The inner, nacreous, layer is pro-
duced only in the anterior portion of the valves and mainly in the area between the two
adductor muscles (Plate 154, figs. 2, 4). It is composed of flat, shingled layers of arago-
nitic, nacreous material which parallel the long axis of the valves. The extent and shape
of the nacreous layer are important characters in the classification of this family as is
shown in the systematic portion of this paper. One of the earliest studies to be made on
the structure of the shell was that of Carpenter (1844) in which he described the irregu-
larly hexagonal crystals and the network of conchiolin which holds them together. Ad-
ditional contributions were made by Biedermann (1901) and Karny (1913). A summary
of this work as well as a brief discussion of the shell structure of Recent and fossil Pin-
nidae and related forms is given by Béggild (1930) in his interesting paper, ‘“The Shell
Structure of the Mollusks.”’

Hinge teeth are lacking and the valves are united along their dorsal margin by a liga-
ment which is divided into two main portions which C. M. Yonge (1953) has termed the
primary and secondary ligaments. The ligament proper (primary ligament) extends from
the anterior adductor muscle posteriorly to the posterior margin of the inner nacreous
layer. ‘The fusion of the valves on the dorsal margin posterior to this is a result of the
fusion of the mantle lobes and, though called the secondary ligament, is not a true liga-
ment. The primary ligament in the Pinnidae is not elastic but it is, however, similar in
structure to that found in other bivalves.

Nores oN ANATOMY AND PHYSIOLOGY

The Pinnidae have a large, more or less centrally located posterior adductor muscle
and a very small anterior adductor muscle. The soft parts are confined mainly to the area
between the two muscles; only the greatly enlarged posterior lobes of the mantle and
the elongate ctenidia or gills extend beyond the posterior adductor muscle. Both the
mantle and the gills are capable of retraction to the muscle. There are no true siphons,
but the mantle cavity is divided into incurrent and excurrent chambers by an intermantle
septum. The anatomy of Pinna nobilis Linné has been described and figured by Poli
(1795), that of Atrina rigida Solander by B. H. Grave (1911) and of Pinna carnea Gmelin
by C. M. Yonge (1953). Since the general morphology of the soft parts of these species
varies but little they will be discussed together, the differences being noted where necessary.

There are many unique characters in this family which fit it for its peculiar mode of
life. Members of the Pinnidae generally live embedded vertically in a soft substratum
attached to stones or other objects by a large byssus. Usually that portion of a shell not
occupied by the soft parts (i.e., the portion posterior to the posterior adductor muscle)
protrudes above the surface where it is exposed and is often broken. This protrusion of
the shell above the soft substratum, however, allows the animal, which does not have ex-
tendible siphons, to get water from well above the bottom and so to avoid much of the
debris which might otherwise be drawn into the mantle cavity. The Pinnidae burrow
more deeply into the substratum as the shell grows, the anterior end being worn away
while the growth of shell is almost entirely at the posterior margin. Though the mantle

288 JOHNSONIA, No. 38 Pinnidae

adheres closely to the shell it is attached only at two points and so can be readily re-
tracted. The greatly enlarged and retractable portion of the mantle is supplied with
numerous pallial muscles which radiate from the two pallial retractor muscles as shown
in Plate 152. The fact that the mantle can be retracted quickly on the slightest stimulus
prevents injury to the mantle proper and thus allows rapid repair of the shell if it has
been broken. When completely retracted the mantle and gills he folded closely against
the posterior adductor muscle and in this condition cover only the area corresponding to
the inner nacreous layer of the shell.

A unique and very important structure in the Pinnidae is the waste canal by means of
which particles of sand and other debris are carried from the incurrent portion of the
mantle cavity. The waste canal consists of slightly raised channels, one on each lobe of
the mantle, which oppose each other to form a tube. The inner surfaces of the channels
are lined with strong ciliaand are supplied with mucous glands. The canal extends nearly
the length of the mantle and parallels the outer margin of the gills. All material enter-
ing the canal is carried rapidly posteriorly (upward) and is consolidated with mucus into
strings and ejected. Although the waste canal is highly developed in the Pinnidae, it is
analogous to the ciliary tracts which function in the ejection of particles from the incur-
rent siphon in other bivalves.

-

ut

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,
ee

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Plate 151. Pinna carnea Gmelin, shell structure. Fig. 1. Section to show the length of the crystals of the
prismatic layer (i.e., thickness of the shell). Fig. 2. Section to show surface of the crystals in cross section
from the same area of the shell as in Fig. 1. (Both 110x.)

Western Atlantic JOHNSONIA, No. 38 289

The mantle is important in cleaning and in repairing the shell, but the Pinnidae pos-
sess in addition a unique structure, the pallial organ, which, according to C. M. Yonge,
probably functions only when the shell has been broken. At such times, after the mantle
has been retracted, the pallial organ may be distended so that it is large and turgid. It
can then be protruded to the edge of the valves where it actively pushes out broken pieces
of shell and debris. The pallial organ at such times is extremely active and many of its
movements appear aimless, but there seems to be little doubt that its function is that of
clearing away the larger particles which are too heavy for the cilia of the mantle to han-
dle. The mantle gradually extends to the edge of the break and the repair and rebuild-
ing of the shell begin. These shells are so fragile and so subject to breakage that one
seldom finds a large specimen which does not show evidence of numerous breaks. Grave
(1911) reported that a hole one half by three fourths of an inch was completely repaired
in three days.

Our observations on living specimens would indicate that the pallial organ functions
whenever the mantle is partially contracted, even if the shell is not broken. If one valve
of Pinna or Atrina is carefully removed and the specimen kept in a pan of well-aerated

;

aT

4
c =>

reproductive So Se ; 8

organs— Ws bs
Sa & : « :
S 3S x %& »
\ ae \ arg
intestine Ss : Ne :
digestive gland SSF Z S :

crystalline style

ventral pallial
retractor

combined mid-gut and style-sac posterior adductor muscle

gastric shield posterior pedal retractor

stomach dorsal pallial retractor

labial palp

anterior pedal retractor

byssus anterior adductor muscle

Plate 152. Semidiagrammatic drawing of the anatomy of Alrina rigida Solander. The right half shows only
the muscles and the characteristic features of the mantle.

290 JOHNSONIA, No. 38 Pinnidae

sea water, the live animal can be observed easily. With the mantle partially retracted
due to the fact that one valve had been removed, the material carried posteriorly by the
waste canal could not be carried beyond the valve and it collected at the posterior end
of the canal. When a sizeable ball of this matter had collected, the pallial organ (in one
of the three live specimens studied) extended and pushed it out of the shell. When not
observing the animal we replaced the valve and in this way were able to keep it fairly
active for three days.

On Plate 152 we present a semidiagrammatic drawing of the gross anatomy of Atrina
rigida Solander. To make this sketch the adductor muscles were cut just inside the
mantle on one side so that only the mantle and portion of the cut muscles were left
adhering to the right valve. On the left valve we have shown the animal as it can be seen
without dissection. We have, however, indicated (as if the animal was transparent) the
esophagus, stomach, combined mid-gut and style sac, and the intestine which cannot be
seen without dissection for these are embedded in the large digestive gland and repro-
ductive organs. The crystalline style in Atria is unusually large—a specimen 150 mm.
long having a style 48 mm. long and 2 mm. in diameter at the anterior end.

Purchon (1957) described in detail the stomachs of Atrina vevillum Born and Pinna
atropurpurea Sowerby, both species of the Indo-Pacific region. He mentioned in par-
ticular the development of a long slender tongue on the major typhlosole, a fold in the
interior of the stomach. This tongue extends into the food-sorting caecum of the stom-
ach, greatly increasing the efficiency of the latter. This development of the tongue on
the typhlosole is shared with the Arcidae, Glycymeridae, Mytilidae, Vulsellidae [=
Isognomonidae }, Pteriidae and Ostreidae. Purchon considered that the evolution of such
a tongue could neither have arisen independently in several lines nor did he believe that the
tongue, once produced, could have been lost independently. Therefore he concluded that
these families are monophyletic, but before definite relationships between these families
can be established, studies of other organs and the shells must be made. Certainly the
Mytilidae, Vulsellidae, Pteriidae and Pinnidae are close in their relationship and this
similarity in stomach structure is one more factor relating them. Our observations on the
stomach of Atrina rigida Solander agree closely with those of Purchon.

A discussion of pinnaglobin, the brown pigment in the body fluid of the Pinnidae, was
given by Suto (1938). Suto did not believe that it had much oxygen-holding capacity
and he suggested that further work is needed on this problem.

In a recent paper on the physiology of Pimna muscle, Abbott and Lowy (1956) have
shown that, in this group, the posterior adductor muscles behave similarly to those of
other bivalves. They show that there is a fast and slow acting portion to the muscle and
that the economy of these muscles is due to their slow rate of relaxation following con-
traction. On this basis it is interesting to note that, though the Pinnidae do not have an
elastic ligament against which the muscles must work to keep the shells closed, the valves
themselves, due to their high organic content are flexible and probably take over the
function of the ligament in opening the valves.

Economic IMPORTANCE

The Pinnidae cannot be classed rightfully among the important economic mollusks
though probably no other family of mollusks yields so many products. The large adduc-
tor muscles of both Pinna and Atrina are eaten in many countries where species of these

Western Atlantic JOHNSONIA, No. 38 29]

genera occur. However, European writers state that they require much cooking and are
rather tough, though of excellent flavor.

In Japan, according to A. R. Cahn (1951), there is an active and important fishing in-
dustry for Atrina japonica Reeve. The posterior adductor muscles are marketed fresh and
the clam meats, exclusive of the muscles, are sold either raw or as dried meats which have
first been boiled. They are highly esteemed as food. Even the discarded shells are used
as a cultch for seed oysters and, when ground, they are used as fertilizer. No part of the
animal is wasted in these Japanese fisheries. When Atrina occur intertidally they are
harvested on the night low tide by torchlight, using a stick with a hook to extricate the
shell. In areas where the species occurs in deeper water, fishing is done by divers operat-
ing from a boat. They use a regulation diving suit with air hose and work for periods as
long as six hours in depths of twenty meters. The clams are usually pulled out with a
barbed spear which penetrates the valves of the shell. So important is this fishery in
Japan that there are government regulations concerning the season for harvesting and
the quantity that can be taken.

On the west coast of Mexico in the vicinity of Mazatlan, Atrina maura Sowerby and
Pinna rugosa Sowerby are important items of food among the natives. Pilsbry and Lowe
(1932) described the heaps of empty shells found there and stated that the large muscle
tastes like that of giant scallops. Mexicans call the shells ‘‘hachas’’ or hatchets and they
are fished in much the same manner as that described for the Japanese fisheries. Mr.
John Fitch of the California Fisheries Laboratory has written us recently that there is
an active fishery for Atrina maura at San Blas, Nayarit, Mexico.

Plate 153. The byssus of the Pinnidae. Fig. 1. Byssus of Atrina rigida Solander from Bunch Beach, Florida
to show the long fine threads which may be used in weaving (about = natural size). Fig. 2. Pinna nobilis
Linné, from Cannes, France, the large Mediterranean species which produces a byssus used in the manu-
facture of garments (about } natural size). Fig. 3. A glove knitted from byssus fibers at Tarento, Italy
(USNM no. 149395; about 2 natural size).

292 JOHNSONIA, No. 38 Pinnidae

No one seems to have used the Pinnidae of the Western Atlantic for food though Mr.
Harvey Bullis of the U.S. Fish and Wildlife Service, Bureau of Commercial Fisheries,
Pascagoula, Mississippi says they are excellent either raw or cooked.

All species in this family produce nacreous pearls, and though they are often irregular
in shape, perfect specimens are quite valuable. Pliny mentioned the practice of diving in
the Mediterranean for Pinna in order to obtain the pearls, and Poli (1795) reported taking
twenty pearls from a single specimen of Pinna nobilis Linné. However, it was apparently
in India where the fishing of Pinnidae for pearls was an important industry among the
ancients. In the records of Alexander’s Indian Expedition it was mentioned that in In-
dian seas these shells were obtained by diving for the sake of the pearls. Grave (1923)
stated that he found pearls in about one fifth of all the specimens of Atrina rigida he
examined for his studies. According to his account the pearls were black, usually spheri-
cal and quite smooth. He said they had been used in the manufacture of brooches and
other articles of jewelry and he could see no reason why they should not be used more
extensively. In an interesting paper on the ‘‘Natural History of the Pearls’’ Hass (1955)
described the structure of pearls in the Pinnidae. These pearls have radially arranged
prisms, a condition quite unlike that of all other pearls. This is a result of the unusual
structure of the shell as discussed previously. Because of the radially arranged prisms and
the large amount of organic material between the prisms, the pearls usually fragment on
drying out and so are rather short-lived. As a result they are considered of relatively little
value at the present time. However, pearls produced in the anterior portion of the man-
tle which lays down laminated, nacreous material would be composed of concentric layers
rather than radial prisms. Such pearls should be durable and valuable.

The most famous product produced by the Pinnidae is the byssus fiber, which is an
extremely fine and soft but strong fiber produced by a gland in the foot of the animal
for the purpose of anchoring the shell. The byssus fiber of some of the larger species in
this family is sufficiently long so that it can be spun and then woven or knitted to make
small garments. It has a beautiful golden bronze sheen and was often combined with silk
when used in making larger garments. Most authorities believe that the use of the byssus

ATRINA POSTERIOR } PINNA

posterior
slope

secondory
ligament
/ A Prismatic
ALA BA j \ layer

prismatic
layer
ventral slope

primary
ligament

posterior adductor
muscle scar’ —
E

\
DORSAL ; 5
; Posterior adductor &
‘ muscle scar—#->

!
'

nacreous layer j
VENTRAL dorsal lobe of E

nocreous layers
ventral lobe of

anterior adductor —
nacreous layer

muscie scar.

septo fy ANTERIOR 3)

Plate 154. Diagrammatic sketch of the valves of Pinna and Atrina to show diagnostic characters. Fig. 1.
External surface of valve of Atrina. Fig. 2. Internal surface of valve of Atrina. Fig. 3. External surface of
valve of Pinna. Fig. 4. Internal surface of valve of Pinna.

Western Atlantic JOHNSONIA, No. 38 293

Plate 155. Pinna rudis Linné from Pointe Noire, Gaboon, French Equatorial Africa. Fig. 1. Internal surface
of valve showing the difference in length of the two lobes of the nacreous layer in a large adult specimen
(about 4 natural size). Fig. 2. External surface of the valve showing the large spines of a typical specimen
(about 4 natural size).

294 JOHNSONIA, No. 38 Pinnidae

as a fiber in making garments probably originated in India near Colchi. This is based on
the fact that the earlier Greek and Roman writers referred to Pinna but did not mention
the use of the byssus before the time of Tertullian (150-222 A.D.). Tarento was the
center of the industry in Italy, and Procopius, who wrote on the Persian Wars about
550 A.D., stated that the five hereditary satraps (governors) of Armenia who received
their insignia from the Roman Emperor were given chlamys (or cloaks) made from lana
pinna (Pinna ‘‘wool,”’ or byssus). Apparently only the ruling classes were allowed to
wear these chlamys. Even today asmall remnant of the former industry remains in Italy
and a few articles such as gloves, hats, shawls and stockings are made mainly for the
tourist trade. According to Simmonds (1879) in *“The Commercial Products of the
Sea,’ the byssus formed an important article of commerce among the Sicilians, for which
purpose considerable numbers of Pinna were annually fished in the Mediterranean from
a depth of 20 to 80 feet. He also said, ‘‘a considerable manufactory is established at
Palermo; the fabrics made are extremely elegant and vie in appearance with the finest
silk. Che best products of this material are, however, said to be made in the Orphan
Hospital of St. Philomel at Lucca.*’ Though the modern gloves and shawls are knitted,
the chlamys, gloves and stockings of the ancients were woven, for knitting was not known
until about 1500 according to Yates (1848). Articles made from Pinna byssus are ex-
tremely strong and durable except that they are readily attacked by moths so that great
care must be taken in their preservation. There are, as a consequence, very few examples
of the early garments in existence. On Plate 153 are shown the cleaned byssus of Atrina
rigida Solander; the shell of Pinna nobilis Linné, the species from which the byssus was
obtained for the Italian industry; and a glove made from byssus fiber at 'Tarento, Italy.

The etymology of the word ‘‘byssus’’ was discussed by van der Feen (1949, pp. 66-71).
It was shown that the word was derived from two very similarly sounding Greek terms,
one denoting depth (of the sea or a river), the other referring to fine vegetable fibers
which were woven into valuable fabrics by the ancients. In 1476 Gaza, through an error
in translation of a section concerning Pinna in Aristotle’s **‘Historia Animalium,”’ origi-
nated the use of the word byssus as it is applied to the holdfast fibers of the Pinnidae,
Mytilidae, Pteriidae, etc. He did this by confusing the Greek word for depth, as used
by Aristotle in describing the ecology of Pinna, with the term for the vegetable fibers
and may have genuinely believed that Aristotle intended the word to be applied to the
holdfast. In 1555, Rondelet perpetuated and popularized this mistake and in this way
the term became established. Gesner in 1558 was the first to oppose the erroneous use
of byssus and criticized Gaza’s work. Since that time various workers have taken a stand
against the use of the word and a few have championed it including Sir d’ Arey Thomp-
son in 1910. Although the word byssus is used universally today in referring to the hold-
fasts of bivalve mollusks it is interesting to reflect upon the uncertain basis for its use and
history of origin.

Because the Pinnidae are such unique bivalves and have intrigued man since early his-
toric times there are a host of common names for this group. In France they are known as
‘jambons’ or ham-shells, because of their shape, and the Mexicans refer to them as ‘hachas’
or hatchets. In England they are known as fan-mussels, fin-shells or sea-wings and in the
United States they are often referred to as pen-shells or sea-pens. The Australians refer
to them as razor backs or razor clams as it is so easy to cut one’s feet on them. This com-
mon name should not be confused with that of Hnsis or Siliqua which are also called
razor clams because of their shape.

Western Atlantic JOHNSONIA, No. 38 295

ASSOCIATES OF THE PINNIDAE

The Pinnidae are host to a number of organisms which live both upon the outside of
the valves and in the mantle cavity. Most of the associated organisms use the shell only
as a point of attachment and support. Perry (1936) reported finding a total of twenty-
five different organisms living in association with Atrina rigida. From a fine shipment
of this species which we received from Harvey Bullis, dredged from 3-4 fathoms off
Tampa Bay, Florida, we removed 28 species of mollusks of which 12 were living, as well
as barnacles, tube worms, hydroids, algae, bryozoa, tunicates and brittle stars.

Some of the organisms associated with the Pinnidae, particularly crustaceans, have an
interesting commensal relationship with them. Aristotle was the earliest writer to men-
tion the interesting association of a crab with Pinna. Chemnitz (1785) fully described
this small crab, Pinnotheres,’ which lives in the mantle cavity of Pinna. The crab was
said to leave the cavity periodically, but always to return when danger threatened, thus
warning the Pinna which would thereupon close its valves protecting both itself and the
Pinnotheres within. This fanciful story of the ‘‘friendship’’ of these two animals was
made famous by Pliny and additional accounts of this relationship have been written by
Poli (1795), Montagu (1803), Johnston (1850), Jeffreys (1863) and others. Apparently,

Plate 156. Holotype specimen of Pinna varicosa Lamarck [ =P. rudis Linné] from Trinidad, Lesser Antilles.
Fig. 1. Outer surface of valve. Fig. 2. Inner surface. Fig. 3. Sketch of the inner surface of the valve to
show arrangement of nacreous layer and the muscle scar. The photographs and drawing by courtesy of Dr.
J. Gaillard and the Muséum d’Histoire Naturelle, Paris (all about 4 natural size). This specimen is typical

of the stunted form of rudis as it occurs in the West Indies.

1 Pinnotheres, from the Greek: Pinna--terein, to guard, i.e., the Pinna guardian.

296 JOHNSONIA, No. 38 Pinnidae

wherever members of the Pinnidae occur there are some small crabs which live with them.
In fact, W. L. Schmitt (1931) stated that ‘‘edible mollusks in all parts of the world are
infested by commensal crabs.”* Dr. Schmitt also stated that shrimp have adopted this
way of life, that one rarely finds more than one adult shrimp or crab in a shell and this
is usually a female. The adult males visit from shell to shell, but the females seldom if
ever voluntarily leave the mantle cavity of their hosts. These crustaceans live in their
sheltered homes sharing the food which is brought into the mantle cavity of the mollusk
during its normal feeding process. They are of no benefit to the bivalve as was supposed
by early writers and according to Christensen and McDermott (1958) they probably
‘ause some damage to the gills. This is certainly true of Pinnotheres ostreum Say which
is associated with Crassostrea virginica Linné. These authors made a complete study of
P. ostreum and gave asummary and bibliography of work done on the life history of Pin-
notheres. We have seen specimens of P. maculatus Say taken from Atrina rigida which

Plate 157. Pinna carnea Gmelin from Port Royal, Cat Island, Bahama Islands. Fig. 1. Outer surface of valve
showing the pronounced keel at the anterior end and the few small spines. Fig. 2. Inner surface of valve
showing the arrangement of the nacreous layer (both about 4 natural size).

Western Atlantic JOHNSONIA, No. 38 297

were collected at Sanibel Island, Florida. A shrimp was observed living in the mantle
cavity of a large Pinna carnea at Bimini, Bahama Islands, by Robert Robertson of the
Museum of Comparative Zodlogy. The shrimp sat on top of the posterior adductor mus-
cle of the Pinna in an exposed position, but when disturbed it moved backwards and
down around the muscle until hidden from view.

NoTreEs ON THE FossiL REcorpD

The fossil record shows that the Pinnidae were reasonably well represented in geologic
time. Although many fossils have been reported, very few whole valves have been figured.
This is probably due to the fact that these fragile shells are not readily preserved as fos-
sils. Usually only fragments representing the prismatic layer remain and there is seldom
any indication of the extent or shape of the nacreous layer. As this portion of the shell
is very important in the classification of the family, few conclusions can be drawn from
the fossil record concerning the evolution of this group. Hyatt (1892) discussed the fos-
sil Pinnidae and their relationship to recent forms. He suggested that when material was
available it would probably be found that in the early stages the nacreous layers of Atrina
and Pinna would be quite similar in outline with the carina and division of the nacreous
layer of Pinna appearing at a later stage. Our observations on very young specimens has
shown this to be true as well as the fact that on the embryonic valves the umbos are not
terminal though in the adult stage they virtually come to take this position.

Hyatt, and later Zittel, recorded the family as having existed since the Devonian.
A trina Gray which is known from the Carboniferous is apparently the oldest of the Recent
genera in the Pinnidae, for the earliest known record of Pinna Linné was from the Jurassic.
Another factor which suggests that Atrina is the oldest genus is the internal structure of
the shells. The nacreous layer of Atrina is undivided, with the muscle scar more or less
medial, while that of Pinna is divided by a longitudinal sulcus and has the muscle sear
displaced toward the dorsal margin. These differences may be indicative of a change
from simple to more complex shell structure. It has been pointed out (cf. Beecher 1898)
that in mollusks the development of more complex shell structure is a sign of aging in a
group. Of course, the situation is always complicated by the possibility of a secondary
redevelopment of more simple characters, but this does not seem to have occurred in
the Pinnidae.

Paleontologists have assigned several extinct genera to the family Pinnidae. One of
these, Palaeopinna Hall 1870 from the Devonian, is the earliest record for the family.
The shell of Palaeopinna (cf. Shimer and Shrock 1944, p. 387, pl. 150, fig. 12) lacks the
external keel of Pinna, has convex umbos, and very fine radiating ribs. Aviculopinna
Meek from the Carboniferous and Permian differs from Recent Pinnidae in that the
umbos are located subterminally so that a small wing is present in front of them. A third
fossil genus, Pinnogena ‘Saussure’ Bronn 1836, was reported from Jurassic and Creta-
ceous rocks. It somewhat resembles an oyster. The figure of Pinnogena seebachi Bohm,
in Zittel (p. 368, fig. 621) shows a nearly entire shell with an enormous elongate, cen-
trally located muscle scar. On the exterior of the valves there are large, convex ribs which
radiate out from a central groove. The genus Sulcatopinna was proposed by Hyatt for
Carboniferous forms having extremely elongate shells with a straight hinge line, approx-
imately terminal umbos and with the valves ridged on the dorsal area. He suggested
that this group may have been the forerunner of Pinna.

298 JOHNSONIA, No. 38 Pinnidae

The general form of a very early member of the Pinnidae gained from a survey of
representative genera is something resembling both Pteria and Pinna. This is most evi-
dent when the fossil genus Aviculopinna is considered. Palaeopinna bears a general re-
semblance to both the Pteriidae and the Mytilidae, families generally placed near the
Pinnidae by neontologists.

There is reason to believe, from the fossil findings, that the family Pinnidae was quite
differently distributed in the past than it is today. Present data show that there are no
species found on the west coast of North America north of Mexico, except Atrina old-
roydi Dall which has been taken in 25 fathoms in San Pedro Bay in southern California.
However, Crickmay (1930, p. 47) reported having found a Pinna in Jurassic strata of
British Columbia, and Vokes (1939, p. 50) described an Eocene Pinna from San Joaquin
Valley, California. Ivanov (1926, p. 175) listed Pinna (Aviculopinna) membranacea Kon.
from the Carboniferous of Moscou [Moscow]. Grant and Gale (1931, pp. 145-147) re-
ported two species of Pinna from the Pliocene and a species of Atrina from the Miocene
of California.

From the foregoing discussions it seems reasonable to conclude (on the basis of shell
structure, anatomy and the fossil record) that the Mytilidae, Isognomonidae, Pteriidae,
Pinnidae and probably the Ostreidae, evolved from some common ancestor.

Novres oN CLASSIFICATION

Though the family Pinnidae is characteristic and easily recognized, classification be-
low the family level is difficult and the opinions of the various authors who have worked
with this family vary considerably. Iredale (1989) recognized seven genera in contrast to
the single genus with three subgenera recognized by Winckworth (1929) and the single
genus with only two subgenera by Thiele (1934). There appear to be, however, three
well marked and easily recognized genera (the subgenera of Winckworth), Pinna Linné,
Atrina Gray and Streptopinna von Martens. This generic classification was used by Habe
(1953) in his paper on the Pinnidae of Japan. He recognized three subgenera of Pinna
and two subgenera of Atrina. Only Pinna s.s. and two subgenera of Atrina are repre-
sented in the Western Atlantic.

On the specific level there is even less agreement among authors. Winckworth in his
catalogue listed 175 names as having been introduced into the literature concerning this
family. Of these he regarded only 29 as being valid species. Iredale commenting on this
great reduction of names said, ‘“‘this seems obviously a reductio ad absurdum, and is
probably as far from the facts as the total number of names is.”’ It should be taken into
consideration, however, that Winckworth included in his catalogue not only all validly
introduced names but also errors in spelling, nomina nuda and names from pre- Linnean
authors and non-binomial works. There are 37 such entries in his catalogue which gives
a ratio of about one in five remaining as a valid species. Just how many species will be
recognized when the Pinnidae of the world have been adequately studied is hard to say
but the number probably will be slightly higher than that of Winckworth, perhaps 45
to 50.

Linné (1758) instituted the genus Pinna and described eight species, three of which are
unrecognizable and a fourth referred to the pen of a Loligo. These have been treated
by Hanley, Winckworth and Dodge. Lamarck also described eight species, the type

Western Atlantic JOHNSONIA, No. 38 299

specimens of which have been discussed by Lamy. The two outstanding illustrated
monographs are those of Chemnitz in the ‘‘Conchylien Cabinet’’ and Reeve in the ‘‘Con-
chologia Iconica.’’ ‘The latter is particularly important as the type specimens of Reeve
and Sowerby (some 39 species) were all figured in this study. Hedley (1924) reviewed
the Australian Pinnidae, recognizing 19 species and two subspecies for that area and
Winckworth covered the species of India and Ceylon. ‘There has been no world-wide
treatment of this group since that of Reeve except the catalogue by Winckworth.

The confused state of the taxonomy of this family and the differences of opinion of the
various authors are probably the result of a number of difficulties inherent in the study of
this group. Shells of the Pinnidae are extremely fragile and have a tendency to fracture
spontaneously on drying out. This fragility, combined with the large size and generally
unattractive appearance of these shells have made collectors reluctant to gather and ship
them. Consequently museum collections are small, they lack comparative material of a
geographic nature and seldom contain lots of more than 4 or 5 specimens from a given

Plate 158. Atrina (Alrina) rigida Solander from Coconut Grove, Florida. Fig. 1. External surface of valve
showing the tubular spines and the sculptured ventral slope. Fig. 2. Internal surface of the valve showing
the nacreous layer and the protruding muscle scar (both about 3/5 natural size).

300 JOHNSONIA, No. 38 Pinnidae

locality. Growth series are almost completely lacking. The Pinnidae as a whole have a
capacity for ecologically induced variation and this has resulted in specimens of species
which are widely separated geographically, such as Atrina inflata Dillwyn of Australia
and Atrina serrata Sowerby of the Western Atlantic, having almost identical external
appearances. However, when series are examined, the two species are found to be quite
distinct. A similar problem may also arise with two species from the same locality, but
actually belonging to different subgenera such as Atrina (Atrina) rigida Dillwyn and
A. (Servatrina) seminuda Lamarck. These factors combined with the lack of large series
in museum collections has resulted in many misidentifications as well as the naming of
numerous forms which were apparently only ecologic variants. If the characteristics of
the inner surface of the valves (the shape and extent of the nacreous layer and the muscle
scars) are considered in conjunction with the external characters, most species can be de-
termined readily. Unfortunately, however, few of the early descriptions either mentioned
or illustrated the interior of the valves. Consequently, when using works such as Reeve’s
‘*Conchologia Iconica,’’ it is often difficult, if not impossible, to place a species in the
proper genus. Winckworth, Hedley and others have mentioned the importance of the
muscle sears and the pattern of the nacreous layer in the classification of this family and
our work has shown that these are among the best taxonomic characters.

Very little is known of the anatomy of most species of the Pinnidae and almost noth-
ing is known of their breeding habits and life history. Further study on these problems
will undoubtedly contribute to an understanding of the classification of the family and
its relationships. Our observations on living animals show that the colors of the various
organs, the position and curvature of the waste canal, as well as the size and shape of the
pallial organ, are all excellent taxonomic characters.

We have included in the synonymies of the various species considered in the systematic
portion of this paper only those names which we were certain referred to the species in
question. Undoubtedly there have been some omissions, particularly in the synonymy
of Pinna rudis Linné, an Eastern Atlantic species which occurs only sporadically in the
West Indies. When reviewing the literature, one finds names such as muricata Linné
applied to species belonging to both Pinna and Atrina, and coming from nearly all parts
of the known range of the family. This is also the case with a number of other names
though to a lesser degree.

ACKNOWLEDGMENTS

Many individuals and institutions have contributed valuable material used in this study
as is shown in the records and we wish to acknowledge all of them. We are grateful to
G. Usticke, St. Croix, Virgin Islands; Germaine Warmke of the Institute of Marine
Biology, University of Puerto Rico, Mayagiiez; Harvey Bullis of the United States
Fisheries Laboratory, Pascagoula, Mississippi; and David and Nevada Schmidt of Sarona,
Wisconsin for the material they collected especially for this study. Particular thanks are
due Jean Foshee and Kathleen Johnstone of Mobile, Alabama and Robert Work of the
Marine Laboratory, University of Miami who have sent both living and preserved ma-
terial. H. A. Rehder of the United States National Museum and R. 'T. Abbott of the
Academy of Natural Sciences, Philadelphia were helpful during our visits to these insti-
tutions to examine the collections. The collections of the California Academy of Sciences
(CAS) and the San Diego Museum of Natural History (SDM) were also examined and

Western Atlantic JOHNSONIA, No. 38 301

we wish to thank L. G. Hertlein and E. P. Chace for their cooperation. For the photo-
graphing of type specimens we are grateful to I. C. J. Galbraith of the Mollusca Section,
British Museum (Natural History) and J. Gaillard of the Muséum d’ Histoire Naturelle,
Paris. Credit for these photographs is given in the plate captions. To W. J. Clench,
M. E. Champion and our associates in the Mollusk Department, we are grateful for the
discussion of several problems and for the reading of the manuscript; R. Robertson in
addition aided in making the sections of the Pinna shell (Plates 149 and 150). We are
indebted to Professor John H. Welsh, Biological Laboratories, Harvard University, for
discussing with us certain physiological problems.

WESTERN ATLANTIC PINNIDAE

The species of Pinnidae in the Western Atlantic may be identified by means of the
following key which is based on adult specimens (see Plate 154).

1. Nacreous layer divided by asuleus ........ 2... . .2 (Pinna)
Nacreous layer not divided by a sulcus. . ... oe es Aiea
2. Ventral lobe of nacreous layer longer than dorsal ibe: valves usually pale
red-orange, thin, with few if any spines; ribs 8-12 . . . . . Pinna carnea
Dorsal lobe of nacreous layer longer than ventral lobe; valves usually dark
red-brown, thick, with large spines; ribs 5-7 (in the Western Atlantic usu-
ally stunted, deformed and devoid of spines, but in Eastern Atlantic large

and spinose) . age A SP ren ‘ End . « «.» « “Pmna-rudis
3. Posterior Rutactor muscle scar rea: Ba ae nacreous layer ..... 4
Posterior adductor muscle scar protruding beyond posterior border of nacreous
AGED (ote 1. be oan aa om. eos) a ss se we Abine rieda
4, Valves with numerous, crowded, fine ae ee only with imbrications; ribs
over 30; shell light greenish tan . ..... oe Se Deine ‘serrata
Valves usually with few ribs armed with spines; aie less than 20; shell gray-
isht Gum tomprteke «60 wee cd es a tele Bovdl's .) o g re, 43>. lina Semin

Genus Pinna Linné

Pinna Linné 1758, Systema Naturae, ed. 10, p. 707 (type species, Pinna rudis Linné, by subsequent selec-

tion, Children 1823, Gray 1847).
Chimaera Poli 1791, Testacea Utriusque Siciliae 1, p. 31 (type species, Pinna nobilis Linné, by subsequent

selection, Winckworth 1929).
Chimaeroderma Poli 1795, Testacea Utriusque Siciliae 2, p. 259 (type species, Pinna nobilis Linné, here

selected).
Pinnarius Duméril 1806, Zoologie Analytique, p. 169, 340; Iredale 1939, Great Barrier Reef Expedition,

Scientific Reports 5, no. 6, p. 309 [new name for Pinna Linné].
Pinnula Rafinesque 1815, Analyse de la Nature, ou Tableau de 1’Univers et des Corps Organises, Palerme,

p. 147 (a substitute name for Pinna Linné).
Cyrtopinna Mirch 1853, Catalogus Conchyliorum Comes de Yoldi, part 2, p. 51 (type species, P. incurva

Gmelin, by monotypy).
Type species, Pinna rudis Linné, by subsequent selection, Children 1823, Gray 1847.
Shell medium to large, reaching about 700 mm. (about 26 inches) in length, wedge-

shaped, generally thin in structure and often sculptured with spinose or imbricate radiating
ribs. Nacreous layer divided by a longitudinal sulcus into a dorsal and a ventral lobe, the

302 JOHNSONIA, No. 38 Pinnidae

large posterior adductor muscle scar being on the dorsal lobe of the nacreous layer adja-
cent to the ligament.

This genus is found in nearly all tropical and subtropical seas with a few species reach-
ing into temperate waters. Specimens are usually found living in sandy-muddy bottoms
where they are firmly attached by large byssi, though in some areas they may be found
living in crevices on rocky shores. A discussion of the biology of the group is given in
the introduction.

There has been a great deal of controversy concerning the type species of this genus.
Since Children’s work (1828) it has been considered as Pinna rudis Linné. However, as
Iredale (1939) has pointed out, Children selected types only for Lamarckian genera and
so, even though Linné’s and Lamarck’s interpretations of Pinna were the same, the first
selection of a type species of Pinna Linné was that of Gray in 1847. He selected as type
Pinnarudis Linné and this selection has been recognized since that time. Unfortunately,
Grant and Gale (1931), attempting an absurdly strict interpretation of the Rules of Zoo-
logical Nomenclature, stated that according to Article 80d and Opinion 16 the type
would have to be Pinna muricata Linné. They based their argument on the fact that

Plate 159. Variations with age in Atrina (Atrina) rigida Solander and Atrina (Servatrina) seminuda Lamarck.
Figs. 1-4. Atrina rigida Solander from off Tampa Bay, Florida in 3-4 fathoms. A series from one locality
showing the gradual protrusion of the posterior adductor muscle scar with increase in age. Figs. 5-8. Atrina
seminuda Lamarck from Aransas Bay, Texas. A series to show the relatively smaller posterior adductor
muscle sear and its inclusion within the nacreous area. All drawings were made with the aid of a Berville
camera lucida.

Western Atlantic JOHNSONIA, No. 38 303

under this species Linné in the Systema Naturae, ed. 10, p. 707, referred to ‘‘Concha
Pinna” in Hasselquist,* making this the type by absolute tautonomy, This has upset a
designation which has been understood for well over one hundred years. In addition,
even Grant and Gale admitted that ‘‘there has been doubt about the recognition of
Pinna muricata Linné.’” Hanley (1855) discussed fully the confusion concerning this
species and Winckworth (1929), following Hanley, stated that the original description
of muricata Linné was a composite, but that Linné inthe Museum Ulricae (1764, p. 545,
no. 143) gave a recognizable description of this species and limited the references to
Rumphius (1741, pl. 46, fig. M) and Gaultieri (1742, pl. 79, fig. D). The locality, ‘‘M.
Mediterraneo,’ as given by Linné in 1758 has long been considered in error as there is
nothing in the Mediterranean which approaches his description. Thus, the name muricata
Linné had been restricted to the Indo-Pacific species and the reference to Hasselquist
eliminated.

Therefore, considering the confusion as to the identity of P. muricata Linné, the sub-
sequent limitation of the name by both Linné and Winckworth, and especially the fact
that the Hasselquist reference was to a species which no one has been able to recognize,
we agree with Iredale (1939) and Habe (1953) that the type species of Pinna Linné
should remain Pinna rudis Linné, Gray 1847.

Plate 160. Atrina(Servatrina) seminuda Lamarck. Holotype specimen of Pinna (Atrina) seminuda Lamarck from
Brasil. Fig. 1. External view of left valve. Fig. 2. Internal view of right valve (both natural size). Photo-
graphs courtesy of the Muséum d’Histoire Naturelle, Paris.

' Hasselquist, F. 1757, Iter Palaestinum, p. 448 [not 447 as given by Linné 1758], no. 137, Stockholm.

,

304 JOHNSONIA, No. 38 Pinnidae

Pinna rudis Linné
Plate 150, figs. 3-4; Plates 155, 156

Pinna rudis Linné 1758, Systema Naturae, ed. 10, p. 707 (O. Meridionali, Mediterraneo); Hanley 1855,
Ipsa Linnaei Conchylia, p. 148; Dodge 1952, Bulletin American Museum of Natural History 100, p. 223.

Chimeroderma rude Poli 1795, Testacea Utriusque Siciliae 2, p. 259.

Pinna ferruginea Réding 1798, Museum Boltenianum, p. 160 (refers to Chemnitz 8, pl. 88, fig. 773).

Pinna elongata Réding 1798, Museum Boltenianum, p. 160 (refers to Chemnitz 8, pl. 88, fig. 773).

Pinna pernula Roding 1798, Museum Boltenianum, p. 160 (refers to Chemnitz 8, pl. 92, fig. 785, St. Croix,

Virgin Islands).
Pinna varicosa Lamarck 1819, Histoire Naturelle des Animaux sans Vertébres 6, p. 133 (I’ile de Ja Trinité

(Trinidad, Lesser Antilles] ).
Pinna paulucciae de Rochebrune 1883, Bulletin de la Société Philomathique de Paris (7) 7, p. 181 (estuary

of Casamance River, Sénégal); Lamy 1911, Bulletin Muséum d’Histoire Naturelle, Paris 17, p. 319.

Description. Shell reaching 565 mm. (about 22 inches) in length, wedge-shaped, sub-
inflated, with a weak to moderately strong longitudinal keel at the anterior end. Valves
thin in young specimens, but reaching a thickness of 3.5 mm. in large adults which are
sculptured with radiating ribs. Shells translucent, particularly in young specimens, and
ranging in color from a red-orange to a dark, reddish brown. Surface generally glossy.
Sculpture consisting of from 5 to 8 strong, radiating ribs which support large tubular
spines which open posteriorly. Concentric sculpture consisting of inconspicuous growth
lines. Posterior margin irregular, but usually evenly rounded, with occasional specimens
being truncate. Dorsal margin usually straight. Ventral margin convex posteriorly, con-
cave anteriorly. Interior of the valves a deep red-orange and glossy. Nacreous layer iri-
descent and extending from the anterior end of the shell to the posterior margin of the
posterior adductor muscle scar. ‘The nacreous layer is divided by a longitudinal sulcus
which extends anteriorly about two-thirds the distance to the anterior adductor muscle
scar. The dorsal lobe of the nacreous layer is usually considerably longer than the ven-
tral lobe, though in young specimens the lobes may be of about the same length. The
posterior margin of the ventral lobe is usually nearly straight, while that of the dorsal
lobe usually has an embayment dorsal to the muscle scar. Anterior adductor muscle scar
small, subapical and nearly as wide as the valve. Posterior adductor muscle sear large,
oval in outline, bounded posteriorly by the end of the nacreous layer and ventrally by
the longitudinal sulcus. Primary hinge ligament thin, black and extending from the an-
terior end of the shell to the posterior border of the nacreous layer. Secondary ligament
not colored but evident in fresh specimens, the dorsal margins of which are fused through-
out: usually in dried specimens the valves gape from the end of the primary ligament to
the posterior margin. Embryonic valves subtriangular, not inflated, the umbos small,
posterior to the center and directed slightly posteriorly (Plate 150, figs. 3-4).

length width
565 mm. 175mm. Pointe Noire, Gaboon, French Equatorial Africa
Ae ete és ec ie st ec “6
300 113 West Africa
192 93 rot ae
161 70 Gorée, Sénégal, French West Africa
139 65 Tobago, Lesser Antilles
202 90 Holotype of Pinna varicosa Lamarck, Trinidad, Lesser Antilles;

measurements according to Lamy.

Western Atlantic JOHNSONIA, No. 38 305

Types. According to Hanley (1855) and Dodge (1952) there is no specimen of Pinna
rudis in the Linnaean collection. Linné’s reference was to d’ Argenville 1742, |’ Histoire
Naturelle, pl. 25, fig. F, which we here select as the type figure. It is a poor figure, but
combined with Linné’s description could certainly refer to this species. Hanley stated
that the name rudis has generally been associated with the figure of Chemnitz (1785) in
the Conchylien-Cabinet 8, fig. 773. The type locality was given by Linné as the Medi-
terranean Sea. The type of Pinna paulucciae de Rochebrune is in the Muséum d’ His-
toire Naturelle, Paris, according to Lamy (1911, p. 319). The type of Pinna varicosa
Lamarck is also in the Paris Museum and the type locality is Trinidad. Lamarck’s type
is a stunted specimen, typical of P. rudis of the West Indies. Through the kindness of
Dr. J. Gaillard of the Muséum d’Histoire Naturelle, Paris, we are able to figure the
holotype specimen (Plate 156). Réding made reference to Chemnitz 8, pl. 88, fig. 778
under both Pinna ferruginea and P. elongata, and this figure is here selected as the type

Plate 161. Alrina (Servatrina) seminuda Lamarck. Fig. 1. External view of the left valve of a specimen from
Peruhybe, Itanhaem, Sao Paulo, Brasil, showing the smooth ventral slope and the open flute-like spines.
Fig. 2. Internal view of right valve of a specimen from Itanhaem, Sao Paulo, Brasil, showing the well-inset
muscle sear (both slightly enlarged).

306 JOHNSONIA, No. 38 Pinnidae

figure for these two names. The type figure, here selected, of P. pernula Réding is Chem-
nitz 8, pl. 92, fig. 785, the only figure to which Réding referred. The type locality of
P. pernulais St. Croix, Virgin Islands, those of the other two Réding names are unknown.

Remarks. See also Remarks under Pinna carnea Gmelin. As indicated in the measure-
ments, Pinna rudis in the Eastern Atlantic is a large, robust species. It is closely related
to carnea Linné and the young are easily confused with that species. However, in P.
rudis, the dorsal lobe of the nacreous layer usually extends well posterior to the ventral
lobe, with occasional specimens, particularly young, having the two lobes of equal length.
In P. carnea the two lobes may be about equal in length, but usually the dorsal lobe is
shorter than the ventral lobe. The ventral lobe of the nacreous layer is usually truncate
in rudis, while in carnea it is rounded to pointed.

We have seen three lots of P. rudis from the Western Atlantic. All are small and de-
formed and they appear to have been living under adverse conditions. There are a num-
ber of records of Pinnidae taken from floating logs, ship bottoms, buoys and other such
objects and the few records of rudis from the West Indies are apparently the result of
such mechanical transport. Though these introduced adults may spawn, conditions for
the growth of the young are apparently unsatisfactory for, to our knowledge, no lasting
colony of this species has ever been established in the West Indies. Specimens of P.
rudis in the West Indies are a much darker red-orange and have a more solid shell than
specimens of P. carnea of the same size.

In the synonymy of Pinna rudis Linné we have included only those names which we
considered were without question synonyms of rudis. There has been so much misiden-
tification and misinterpretation of rudis Linné as well as other species in the Eastern
Atlantic that if we included a reference to each of the misidentifications, the synonymy
would become meaningless.

Range. EasteERN ATLANTIC: Western Mediterranean, Madeira, Canary and Cape
Verde Islands, and the Atlantic coast of Africa south to St. Paul-de-Loanda, Angola.

WestrERN ATLANTIC: Occurs sporadically in Puerto Rico and the Virgin Islands south
to Trinidad and Tobago, Lesser Antilles.

Specimens examined. EastERN ATLANTIC: CANARY IsLaNps: Ténériffe (MCZ).
Frencu West Arrica: Gorée, Sénégal (A. Humes). FreNcH EquaroriaL AFRICA:
Pointe Noire, Gaboon (A. Humes). BELGIAN Conco: Banana (SDM). ANcGo.ta: St.
Paul-de-Loanda (MCZ).

WeEsTERN ATLANTIC: PuERTo Rico: Aguadilla (A. Phares). VirGin Is_Lanps:
Christiansted, St. Croix (G. Usticke). LEssEr ANTILLES: Tobago (MCZ); Trinidad
(Muséum d’ Histoire Naturelle, Paris).

Pinna carnea Gmelin
Plates 149: 150, hes. 1-2-1510. St57

Pinna carnea Gmelin 1791, Systema Naturae, ed. 13, 1, p. 3365 (no locality given) {refers to Knorr 1771,
Verlustiging 2, pl. 23, fig. 1].

Pinna degenera Link 1807, Beschreibung der Naturalien-Sammlung der Universitit zu Rostock, p. 159 (no
locality given) [refers to Chemnitz 8, pl. 87 [not 86 as given in Link], fig. 769.

Western Atlantic JOHNSONIA, No. 38 307

Pinna flabellum Lamarck 1819, Histoire Naturelle des Animaux sans Vertébres 6, p. 130 [in part, references
to Chemnitz 8, pl. 86 [87], fig. 769, and to Pinna carnea Gmelin].

Description. Shell reaching 270 mm. (about 11 inches) in length, wedge-shaped, sub-
inflated, with a weak to moderately strong longitudinal keel at the anterior end; valves
thin, fragile, and sculptured with radiating ribs. Shell translucent, particularly in young
specimens, and ranging in color from light tan to pink and salmon with occasional speci-
mens becoming rather dark orange-brown. Surface generally glossy (except where in-
crusted with organisms and mud). Sculpture consisting of from 8 to 12 moderate to
strong radiating ribs, often with intermediate riblets. These ribs may be smooth or pro-
vided with a few hollow spines which open posteriorly. Concentric sculpture consisting
of inconspicuous lines of growth which are convex posteriorly. Posterior margin irregu-
lar, usually undulate and frequently fractured; it is convex, oblique or nearly straight.
Dorsal margin variable, ranging from broadly concave to nearly straight. Ventral mar-
gin convex to slightly concave, but usually convex posteriorly and concave anteriorly.
In some specimens it may be nearly straight. Interior of valves glossy and light tan to

Plate 162. Holotype of Pinna listeri d’Orbigny |= Altrina seminuda Lamarck] from Brasil (about 3/5 natural
size). Photograph courtesy of the British Museum (Natural History).

308 JOHNSONIA, No. 38 Pinnidae

salmon or orange-brown in color. Nacreous layer iridescent and extending from the an-
terior end of the shell to and including the area of the posterior adductor muscle sear. It
is divided by a longitudinal sulcus which extends nearly to the anterior end of the valve.
Ventral lobe of the nacreous layer usually rounded and longer than the dorsal lobe though
in young specimens the two lobes are often approximately equal in length. Anterior
adductor muscle scar small, nearly as wide as the valve and located just anterior to the
termination of the longitudinal sulcus. Posterior adductor muscle scar large, bounded
posteriorly by the end of the nacreous layer and ventrally by the longitudinal sulcus.
Primary hinge ligament thin, black and extending from the anterior end of the shell to
the posterior border of the nacreous layer. Secondary ligament not colored but evident
in fresh specimens, the dorsal margins of which are fused throughout; not usually seen
in dried specimens the valves of which often gape from the end of the primary ligament
to the posterior margin. Embryonic valves triangular, not inflated, the umbos central,
prominent and directed medially (Plate 150, figs. 1-2).

length width
270 mm. 122mm. Bermuda
258 121 Alice Town, N. Bimini Id., Bahama Islands
Q47 108 Fish Point, Guantanamo Bay, Cuba
192 99 West Summerland Key, Florida
92 5a St. Thomas, Virgin Islands

Types. Gmelin’s only reference under Pinna carnea was to Knorr 1771, pl. 28, fig. 1,
and this is the type figure. The type locality is here designated as Guantanamo Bay,
Cuba, an area from which we have seen material and from which specimens could have
reached Europe at that early date.

Remarks. There has been considerable discussion concerning the species of Pinna in
the Western Atlantic. This has arisen from the fact that species in this genus are vari-
able, that carnea Gmelin of the Western Atlantic and rudis Linné of the Eastern Atlan-
tic are close in their relationship and that young or worn specimens are difficult or even
impossible to distinguish. When reviewing the literature for this group it was found that
about half of the authors referred to all Western Atlantic specimens as carnea while
others considered them to be rudis; still others have considered that both species are
found here side by side. We agree with Dodge (1952) and Abbott (1954) that the com-
mon species throughout the West Indies is Pinna carnea Gmelin, though rudis does
occur rarely in Puerto Rico and the Lesser Antilles. Though the West Indian specimens
of rudis are small and usually lack spines they are readily distinguished by the pattern
of the nacreous layer and muscle scar. Pinna rudis Linné in the Eastern Atlantic is a
large, heavy species, having only 5-8 radiating ribs and much coarser spines than those
of carnea. Specimens of Pinna rudis may reach 565 mm. in length and have spines 10.7
mm. wide at the base. The largest specimen of carnea in the collection of the Museum
of Comparative Zodlogy is only 270 mm. in length with spines 4.5 mm. wide at the
base. See Remarks under P. rudis Linné.

An interesting account of the anatomy and physiology of Pinna carnea is given by
C. M. Yonge (1953). Structures peculiar to the Pinnidae in general are discussed and
comparisons are made with other groups regarding form and evolution.

Western Atlantic JOHNSONIA, No. 38 309

We have seen specimens of this species taken from a buoy off Key West, Florida and
from the inside of an ocean buoy anchored just south of Miami. These specimens were
all small (averaging about 50 mm. in length), thin, and medium to pale salmon in color.
A single specimen taken from a buoy block from off Key West had reached a length of
180 mm. and was a normal P. carnea even though it had lived entirely on the surface of
the block with no opportunity to burrow. This species, occurring as it does on buoys, must
have a free swimming larval stage.

Range. From Lake Worth, Florida, south through the Florida Keys: Bermuda; the
West Indies and from Veracruz, Mexico south probably to Trinidad.

Specimens examined. FLoripa: South Inlet, Lake Worth (MCZ; USNM): Crandon
Park, Miami; off Miami from the inside of an ocean buoy (both R. Merrill); Biscayne
Key (J. K. Howard; J. Schwengel); 4 miles NE of The Elbow, Key Largo in 66 fath-
oms; off Carysfort Light, Key Largo (both MCZ); reef off Teatable Key (Peabody
Mus., Yale Univ.; J. Schwengel; ANSP); Tavernier Key (R. Flipse); Little Duck

Plate 163. Holotype of Pinna subviridis Reeve [ = Atrina seminuda Lamarck | from South Carolina (about + nat-
ural size). Photograph courtesy of the British Museum (Natural History).

310 JOHNSONIA, No. 38 Pinnidae

Key (MCZ; G. Kline); Grassy Key (MCZ; ANSP; G. Kline); Pigeon Key; Missouri
Key; Ohio Key; West Summerland Key (all D. and N. Schmidt); Key Vaca (J.
Schwengel:; D. and N. Schmidt); Big Pine Key (MCZ); Looe Key (J. Schwengel);
Key West (MCZ; ANSP; USNM; CAS); off Key West from a buoy (R. Merrill);
Fort Jefferson, Dry Tortugas (MCZ); Bird Key and Garden Key, Dry Tortugas (both
USNM). Bermupa: Richardson Inlet, St. George Island (MCZ); Castle Harbour in
4-5 fathoms (R. Foster). Banama IsLAaNps: West End, Grand Bahama (MCZ); Hope
Town, Great Abaco (G. Kline); south end, Elbow Cay, Great Abaco (R. Robertson);
Alice Town, North Bimini Id. (MCZ; ANSP); North Cat Cay, Bimini Islands (J.
Schwengel): South Riding Rock, 30 miles S of Bimini Island; NW coast South Bimini
(both R. Robertson); Nassau, New Providence (Peabody Mus., Yale Univ.; USNM);
Adelaide Beach, New Providence (R. Robertson); Sandy Point, Savannah Sound, Eleu-
thera (ANSP; MCZ); Port Royal; Arthurs Town and Northeast Point, all Cat Island;
Little San Salvador, 18 miles W of Cat Island; Matthew Town, Great Inagua (all
MCZ); Big Wood Cay, Andros; George Town, Exuma (both G. Kline); Hog Cay,
Exuma (R. Robertson). Cusa: Cabo Cajon, Pinar del Rio; Cayo Jutia and Bahia de
Cabanas, Pinar del Rio (all MCZ); Cayo Francés, Caibarién, Las Villas (P. J. Bermu-
dez; R. Humes); Castillo de Jagua, Bahia de Cienfuegos, Las Villas (ANSP); Caleton
de Don Bruno, 4 miles SW of Cienfuegos and Punta de la Milpa, Cienfuegos, both Las
Villas (both MCZ); Cayo off Punta San Juan, Bahia de los Perros, Camagiiey (ANSP);
Cable Beach and Fish Point, Guantanamo Bay, Oriente (both MCZ). Jamaica: Port
Royal (USNM). Hispantoua: South of Punta Mangle, Bahia de Samana, Dominican
Republic (USNM). Purrro Rico: Aguadilla and Mayagiiez (both A. Phares); Punta
Algorrobo and Punta Guanajibo (both G. Warmke); Culebra Island (USNM). Viren
IsLANDs: near Christiansted, St. Croix (G. Usticke); St. Thomas (MCZ; USNM; San
Diego Mus.); St. John (ANSP; USNM); Tortola and Guana Island (both MCZ);
Gorda Sound, Virgin Gorda (R. Foster). Lesser ANTILLES: Saint Martin (ANSP);
Antigua (ANSP; USNM); Barbados (MCZ; Peabody Mus., Yale Univ.; USNM;
ANSP); Tobago (MCZ). Carippean IsLanps: Old Providence Island (USNM).
Mexico: Veracruz, Veracruz (T. Pulley; USNM). Cotompia: Cartagena (CAS;
SDM); Santa Marta (USNM).

Genus Atrina Gray

Atrina Gray 1840, Synopsis Contents of the British Museum, ed. 42, p. 151 [nomen nudum]; Gray 1842,
Synopsis Contents of the British Museum, ed. 44, p. 83 [described but no species listed]; Gray 1847, Pro-
ceedings Zoological Society of London, p. 199.

Pennaria ‘Browne’ Mérch 1853, Catalogus Conchyliorum Comes de Yoldi, part 2, p. 51 (type species, P.
rigida Solander { = Atrina rigida Solander| here selected), non Pennaria Oken 1815, non de Blainville 1818.

Pinnaria ‘Browne’ Paetel 1890, Catalog der Conchylien-Sammlung, p. 208 [error for Pennaria Browne |.

Type species, Pinna nygra ‘Chemnitz’ Dillwyn, by subsequent selection, Gray 1847.

Shell reaching about 350 mm. (14 inches) in length, wedge- or wing-shaped in outline.
Thin to rather heavy in structure with terminal umbos and often sculptured with imbri-
cated radiating ribs. Nacreous layer extending about two thirds to three fourths the
length of the shell and not divided by a longitudinal sulcus. The large posterior adduc-
tor muscle scar located subcentrally.

Western Atlantic JOHNSONIA, No. 38 311

Species in this genus are found in nearly all tropical and subtropical seas with a few
species reaching warmer temperate waters.

Subgenus Atrina Gray

The external features of the species included in this subgenus are similar to those of
the genus as a whole. The characteristic feature of the subgenus is the protrusion of the
posterior adductor muscle scar beyond the posterior margin of the nacreous area (Plate
159, figs. 1-4).

There is only one species in this subgenus in the Western Atlantic, Atrina rigida
Solander.

Plate 164. Atrina (Servatrina) seminuda Lamarck from Bogue Island, North Carolina (about } natural size).
A typical specimen showing the smooth ventral slope (fig. 1) and the thin, translucent quality of the shell

often evident in this species (fig. 2).

312 JOHNSONIA, No. 38 Pinnidae

Atrina (Atrina) rigida Solander
Plates 158; 159, figs. 1—4
Pinna rigida Solander 1786, Catalogue of the Portland Museum, p. 136, species 3040 (no locality given)
[refers to Knorr 1771, pl. 26, fig. 1].
Pinna carolinensis Hanley 1858, Proceedings Zoological Society London, p. 225 (Carolina Amer. Sept.);
Reeve 1848, Conchologia Iconica 11, Pinna, pl. 34, fig. 66.

Description. Shell large, reaching about 286 mm. (about 114 inches) in length, wedge-
shaped, usually somewhat inflated, thin, fragile and sculptured with spinose radiating
ribs. Shell translucent, ranging in color from a purplish brown to nearly black, but all
specimens when viewed with transmitted light are a mottled golden to rich red-brown.
Surface generally dull, particularly in older specimens. Sculpture variable, ranging from
specimens which are nearly smooth to those having numerous ribs. The sculpture on the
posterior slope consists of from 6 to 17 ribs which usually bear large, slightly recurved,
tubular spines which may reach 15 mm. in length. Young specimens may have only 3 to
4 ribs: the additional ribs are intercalated as the shell grows in size. Sculpture on the
ventral slope variable, ranging from rather heavy spines to fine imbrications. Concentric
sculpture consisting of irregular lines of growth. Posterior margin irregular, usually
rounded but frequently fractured. Dorsal margin usually slightly concave. Ventral mar-
gin strongly convex posteriorly and concave anteriorly. Interior of the valves iridescent,
somewhat mottled, dark greenish brown to purple-black, usually greenish around the
edges. Nacreous layer extending about two thirds the length of the shell. The large
nearly circular posterior adductor muscle scar is situated at the posterior end of the na-
creous layer, its posterior margin protruding beyond it. The extent of protrusion varies
and increases with the age of the shell. Anterior adductor muscle scar small and nearly
as wide as the anterior end of the shell.

In living specimens the posterior, thickened margin of the mantle is black with small,
irregular white vermiculations; the ventral margin is a gray-white. Inside this thickened
margin the mantle is a bright golden orange. It is most intense near the margin, fading
to a pale orange at the waste canal. The ctenidia are a medium to pale translucent
orange-buff. The pallial organ, even when contracted, has a large conical *“‘head’’ which
is an opaque cream color. The stalk of the pallial organ is a translucent grayish ivory.
The foot is cream colored. The sexes are separate and are readily distinguished — the
testes being a cream-white and the ovaries a medium magenta.

length width
286 mm. 180mm. Gulfport, Florida
237 133 Beaufort, North Carolina
235 128 Cayo Francés, Caibarién, Cuba
230 144 Off Pan American Airport, Coconut Grove, Florida
226 158 Alice Town, North Bimini Id., Bahama Islands
215 LM Chadwick Beach, Lemon Bay, Florida
LV 97 Santa Barbara de Samana, Dominican Republic

Types. The location of the type specimens of Pinna rigida Solander is unknown to
us. Solander referred to Knorr 1771, pl. 26, fig. 1, which we here consider the type fig-
ure. The type of Pinna carolinensis Hanley may be in the British Museum (Natural
History), though Wilkins (19538) does not list it. The type locality is here restricted to
Beaufort, North Carolina.

Western Atlantic JOHNSONIA, No. 38 313

Remarks. This isacommon species in the Florida region. It is close in appearance to
Atrina seminuda Lamarck with which it has generally been confused. Externally, the
two species often cannot be separated, but the relative positions of the muscle scar and
the nacreous layer readily differentiate them. In Aftrina rigida the posterior adductor
muscle scar is proportionately larger in relation to the size of the shell. The posterior
margin of the muscle scar is continuous with the posterior border of the nacreous area,
and in older specimens, protrudes well beyond it (Plate 159, figs. 1-4). In Atrina semi-
nuda Lamarck the muscle scar is set within the nacreous area with a border of nacreous
material margining it posteriorly (Plate 159, figs. 5-8). Other characters used in separat-
ing rigida and seminuda are often less definite. However, A. rigida generally has a
heavier shell, which is rounded rather than truncate posteriorly and its dorsal border is
often slightly concave rather than straight as in seminuda. Atrina rigida is usually more
spinose, particularly on the ventral slope which is generally smooth in A. seminuda. In
color, the shell of A. 77gida is far darker, usually being a purple-black in adult specimens,
while in seminuda it is mottled tan and purple. If living material is available 77gida may
be readily separated from seminuda by the color of the soft parts. The most conspicuous
difference is the bright golden-orange mantle coloration of 7igida while the same area is

Plate 165. Atrina (Servatrina) seminuda Lamarck from Port Isabel, Texas. Fig. 1. Outer surface of a typical
specimen from the Texas area showing the nearly smooth, almost spineless valves and the truncate posterior
margin. Fig. 2. Inner surface showing the dark color of the shells from this area (both about 7 natural size).

314 JOHNSONIA, No. 38 Pinnidae

a pale yellow in seminuda. The gills of rigida are a pale orange-buff while those of semi-
nuda are intense brownish orange. The ‘‘head”’ of the pallial organ of 7igida is large and
conical while that of seminuda is small and rounded. See Remarks under A. seminuda
Lamarck.

An account of the anatomy and physiology of Atrina rigida is given by Grave (1911).
This species is usually found living in sandy mud from the intertidal area to depths of
about three to fifteen fathoms. Specimens are very abundant in certain areas and may
be a real hazard to fishermen who report that their nets are often badly torn when drag-
ging over a bed of Atrina.

Atrina tuberculosa Sowerby is the Eastern Pacific analogue of this species, though it
is a larger and heavier shell. It is known mainly from the Gulf of California, ranging from
Puerto Pefiasco, Sonora to La Paz, Baja California, Mexico.

Range. From Cape Hatteras, North Carolina, south through the Florida Keys and
the west coast of Florida as far north as Panama City; the Bahamas; Cuba and His-
paniola. his species apparently does not occur on the northern Gulf coast, in Texas or
in Central America.

Specimens examined. Norru Carouina: Buxton, Cape Hatteras (USNM); Beaufort
(MCZ;: San Diego Mus.); Wilmington (MCZ); Smith Island, Cape Fear (ANSP).
FLoripa: Jacksonville Beach; Daytona Beach; New Smyrna; Cape Canaveral (all
MCZ); Lake Worth (G. Kline; MCZ; ANSP; USNM;; San Diego Mus.); off Hills-
boro Inlet, Pompano (MCZ); Crandon Park, Miami (R. Merrill); Biscayne Bay
(ANSP; R. Work); Key Biscayne, Miami(J.K. Howard); Little Pine Key and Grassy
Key (both ANSP); Sandy Key, Cape Sable (USNM); off Everglades (MCZ); Cape
Romano (ANSP); Bonita Beach, Bonita Springs (MCZ); Punta Rassa (D. and N.
Schmidt); Sanibel Island (MCZ; ANSP; J. S. Schwengel; San Diego Mus.); Captiva
Island (MCZ; J. S. Schwengel); Boca Grande, Gasparilla Id. (ANSP); Chadwick
Beach, Lemon Bay (MCZ); Long Key, off Sarasota (MCZ; USNM); Mullet Key,
Tampa Bay; Boca Ciega Bay, St. Petersburg (both ANSP); St. Petersburg (USNM;
San Diego Mus.); off Tampa Bay in 3-4 fathoms (H. Bullis); Gulfport (MCZ); Cedar
Keys (USNM; CAS; SDM); near Sea Horse Key, Cedar Keys (MCZ; ANSP); St.
Joseph Bay (ANSP); Panama City (USNM). Banama IsLanps: Rocky Point, East
Bimini (R. Robertson); Alice Town, North Bimini Id. (MCZ). Cuspa: Cayo Francés,
Caibarién, Las Villas; Fish Point, Guantanamo Bay (both MCZ); Cayo Galindo, Car-
denas, Matanzas (ANSP). Hispanroia: Santa Barbara de Samana, Dominican Repub-
lic (MCZ).

Subgenus Servatrina Iredale
Servatrina Iredale 1939. Great Barrier Reef Expedition, Scientific Reports 5, no. 6, p. 317.
Type species, Pinna assimilis Reeve, by original designation.
Externally the shells are similar to those of other Atrina. The characteristic feature
of the subgenus is found on the inner surface of the valves. The large posterior adductor

muscle scar is enclosed completely within the nacreous area rather than protruding be-
yond it as in Atrina s.s.

Western Atlantic JOHNSONIA, No. 38 315

The subgenus is world-wide in distribution in the warmer temperate and tropical seas.

Atrina (Servatrina) seminuda Lamarck
Plates 150, figs. 5-6; 159, figs. 5-8; 160-169

Pinna semi-nuda Lamarck 1819, Animaux sans Vertébres 6, pt. 1, p. 131 (les Mers d’Amérique); Lamy
1932, Bull. Muséum National d’Histoire Naturelle (2) 4, p. 896 (Brasil); non P. seminuda Bowdich 1825;
non seminuda “Lamarck” Reeve 1858.

Pinna alta Sowerby 1835, Proceedings Zoological Society London, p. 84 (Sinu Honduras); Reeve 1858,
Conchologia Iconica 11, Pinna, pl. 6, fig. 11.

Pinna listeri d’Orbigny 1846, Voyage l’Amérique Méridionale 5, pt. 3, Mollusques, p. 641, pl. 85, fig. 1
(cdtes du Brésil); Carcelles 1944, Revista del Museo de la Plata (n.s.) Zoologia 3, p. 278, pl. 10, fig. 77.

Pinna patagonica d’Orbigny 1846, Voyage ]’Amérique Méridionale 5, pt. 3, Mollusques, p. 641, pl. 85,
fic. 2 (l’embouchure du Rio Negro, Patagonie); Carcelles 1944, Revista del Museo de la Plata (n.s.) Zoolo-
gia 3, p. 278, pl. 10, fig. 78.

Pinna subviridis Reeve 1858, Conchologia Iconica 11, Pinna, pl. 17, fig. 32 (South Carolina).

Pinna ramulosa Reeve 1858, Conchologia Iconica 11, Pinna, pl. 28, fig. 52 (West Indies).

Pinna d’orbignyi ‘Hanley’ Reeve, Aug. 1858, Conchologia Iconica 11, Pinna, pl. 26, fig. 49 (West Indies);
Hanley, Jan. [Nov.] 1858, Proceedings Zoological Society London, p. 228.

Plate 166. Holotype of Pinna ramulosa Reeve | = Atrina seminuda Lamarck] from the West Indies (about 4
natural size). Photograph courtesy of the British Museum (Natural History).

316 JOHNSONIA, No. 38 Pinnidae

Description. Shell reaching 243 mm. (about 92 inches) in length, wedge-shaped, usu-
ally somewhat inflated, thin, fragile and sculptured with spinose radiating ribs, but with
the ventral slope usually smooth. Shell translucent, ranging in color from a uniform
grayish tan to a mottled tan and purple or purple-brown, particularly when viewed with
transmitted light. Surface usually somewhat glistening, particularly in young specimens.
Sculpture variable, ranging from specimens which are smooth or nearly so to those having
numerous, spinose, radiating ribs. The sculpture on the posterior slope consists of from
5 to 16 radiating ribs which may be smooth or armed with spines. ‘These spines are usu-
ally tubular and slightly recurved, though on occasional specimens they appear as coarse,
upright flutes. Sculpture on the ventral slope usually consisting of fine growth lines and
indistinct radiating ribs, with occasional specimens having small spines near the posterior
margin. Concentric sculpture consisting of irregular growth lines. Posterior margin
irregular, usually somewhat truncated with a distinct angle at the union of the ventral
slope and the posterior slope. Dorsal margin slightly concave. Ventral margin convex
posteriorly and only slightly concave anteriorly. Interior of valves iridescent, ranging in
color from a nearly uniform tan to a mottled tan and purple-brown. Nacreous layer ex-
tending about one half to two thirds the length of the shell. Posterior adductor muscle
scar small in relation to the size of the shell, oval in outline and usually lying well within
the nacreous area. In occasional specimens there is only a very narrow border of nacreous
material edging the muscle scar posteriorly, but the muscle scar never protrudes beyond
the nacreous area. Anterior adductor muscle scar small and nearly as wide as the anterior
end of the shell. Embryonic valves subcircular, inflated, the umbos prominent and di-
rected slightly posteriorly (Plate 150, figs. 5-6).

Plate 167. Alrina (Servatrina) seminuda Lamarck from Kingston, Jamaica. A specimen typical of this species
in the West Indies, showing a slight broadening of the valves posteriorly and an increase in the number
of radiating ribs and spines (about 4 natural size).

Western Atlantic JOHNSONIA, No. 38 317

In living specimens the posterior and thickened margin of the mantle is black with
reddish markings. Inside the margin the mantle is pale yellow, the color being most in-
tense near the thickened margin, fading to white at the waste canal. The ctenidia are an
intense brownish orange and very conspicuous. The pallial organ, when contracted, has
a small rounded ‘“‘head’* which is a light salmon in color. The stalk of the pallial organ
is a translucent white. The foot is light yellow. The sexes are separate—the ovaries are
a bright pink-orange. A male specimen was not seen.

length width
243 mm. 125mm. Charleston, South Carolina
LNT ar 90 Aransas Bay, Texas
169 112 Kingston, Jamaica
151 92 West Trinidad
151 87 Gulfport, Florida
140 67 Tuxpan, Veracruz, Mexico
125 52 Itanhaem, Estado de Sao Paulo, Brasil

Types. The holotype of Pinna seminuda Lamarck is in the Muséum d’ Histoire Nat-
urelle, Paris. The type locality is Brasil according to Lamy (1982, p. 396). Through the
kindness of J. Gaillard of the Paris Museum we are able to figure the holotype (Plate
160). The type locality is here restricted to Rio de Janeiro, Brasil.

The holotypes of the following species are all in the British Museum (Natural History):
P. listeri d’Orbigny from Brasil, no. 1854.12.4.814; P. alta Sowerby from the Bay of
Honduras, no. 1952.8.29.1; P. ramulosa Reeve from the West Indies, no. 1952.8.29.45;
P. subviridis Reeve from South Carolina, no. 1952.8.29.62; and P. dorbignyi Reeve

Plate 168. Paratype of Pinna alta Sowerby | = Alrina seminuda Lamarck] from the Bay of Honduras (about 4
natural size). An unusually broad form. Photograph courtesy of the British Museum (Natural History ).

318 JOHNSONIA, No. 38 Pinnidae

from the West Indies, no. 1952.8.29.15. Through the kindness of I. C. J. Galbraith of
the Mollusca Section of the British Museum (Natural History) we are able to figure all
of the above type specimens.

Remarks. Atrina seminuda Lamarck is a very variable species which has generally
been considered a synonym of A. rigida Solander. However, on the basis of its muscle
scars and the extent of the nacreous layer, it belongs in the same subgenus as JA. serrata
Sowerby. See Remarks under A. serrata. Externally, A. seminuda is often difficult to
distinguish from A. rigida, and for this reason has long been confused with that species.
There is no single, external character which will separate these two species for the range
in variation of each overlaps with that of the other. In general, seminuda is lighter in
color and less strongly sculptured, the ventral slope is usually devoid of spines and the
valves are usually truncate posteriorly. The tremendous range of variation in the sculp-
ture of seminuda appears to be largely a reflection of ecologic conditions. However, on
the basis of the material which we have studied, there does seem to be an increase in the
number of ribs and the size of the spines on specimens from near the center of the range.
Specimens from Brasil and North Carolina are quite similar in appearance, while those
from the West Indies are usually more spinose. Specimens from the Texas coast are
generally darker in color, smoother, often being completely spineless, and more truncate
than those from elsewhere in the range. The relationship of the width to the length of
the shell in this species is also extremely variable, as is shown by the measurements and
illustrations. Atrina alta Sowerby is an example of an unusually broad and spinose form
while d’orbignyi Reeve is a young, elongate and spinose form. Wilkins (1958) believed
the type locality of d’orbignyi Reeve to be in error and placed this species in the synony-
my of sguamifera Sowerby from South Africa. However, there are in the Museum of
Comparative Zodlogy several specimens from the West Indian region which approximate
the figure of the type specimen very closely (Plate 169). We therefore consider d’orbig-
nyt Reeve to be a synonym of seminuda. The most dependable character to use in dis-
tinguishing seminuda from rigida is that of the position, size and relationship of the mus-
cle scar and the nacreous layer. In Plate 159 we have indicated the change in position
of the muscle scar with the growth of the shell for these two species. Young specimens
and those which have grown rapidly are sometimes difficult to distinguish. In some
fairly large specimens of seminuda the muscle scar is only slightly inset in the nacreous
area while in some equally large but rapidly grown specimens of rigida the muscle scar
has not protruded. However, in these cases, other characters, such as the general outline
of the nacreous area, the shape of the shell and the lack of sculpture on the ventral slope
aid in determining the species. The muscle scar of seminuda never protrudes beyond the
nacreous area and it is usually much smaller in proportion to the size of the shell than
that of rxgida. If aseries of specimens from a given locality is available there is no diffi-
culty in determining the species and when working with animals that are so readily af-
fected by their environment, as are all species of Atrina, large series are essential. When
living material is available seminuda is readily differentiated by the coloration of the soft
parts. See Remarks under rigida.

Atrina maura Sowerby is the Eastern Pacific analogue of this species. It is the most
common species in the Eastern Pacific, ranging from Bahia Magdalena, Baja California,
Mexico, south to Peru. This species is commonly eaten by the Mexicans.

Western Atlantic JOHNSONIA, No. 38 319

Range. Weare unable to account for the absence of A. seminuda in southern Florida,
the Florida Keys and the Bahama Islands. Except for these areas, its range compares
with that of Laevicardium laevigatum Linné, Trachycardium muricatum Linné and
Macrocallista maculata Linné all of which extend from North Carolina to southern Brasil
or Argentina.

Specimens examined. NortH CaroLina: Cape Lookout (USNM); Beaufort (ANSP:
USNM): Bogue Id. (MCZ); Long Beach near Southport (USNM): Smith Id., Cape
Fear (ANSP). Sourn Caroiina: 4 miles SW of Myrtle Beach (ANSP); Isle of Palms
(MCZ); Sullivans Id. (CAS; MCZ); Charleston (MCZ). Grorcia: Sea Id. Beach
(ANSP); St. Simons Id. (USNM); Cumberland Id. (MCZ). FrLortpa: Amelia Id.
(ANSP; USNM); St. Augustine (USNM); Anastasia Id., St. Augustine; Daytona
Beach: Lake Worth (all ANSP); Coronado Beach (MCZ); Fort Myers Beach (ANSP):
Sanibel Id. (J. S. Schwengel; ANSP); Captiva Id., and Indian Beach, Sarasota (both
MCZ); St. Petersburg (USNM); Pass-a-grille, Pinellas Co. (MCZ); Clearwater Bay,
Pinellas Co. (ANSP): Gulfport (SDM; MCZ); Panama City (USNM). ALaBama:
Dauphin Island (J. Foshee). Loutstana: American Bay, St. Bernard Parish (R. Parker);
Grand Isle(MCZ); Freemason Id., Chandeleur Islands (D. Moore). Texas: Galveston

Plate 169. Holotype of Pinna d’orbignyi Reeve [= Atrina seminuda Lamarck] from the West Indies (natural
size). Fig. 1. A young specimen which was probably growing in quiet water allowing unusual development of
thespines. Fig. 2. Inner surface of the opposite valve which, though somewhat overgrown with Bryozoa, does
show the muscle sear and nacreous layer. Photographs courtesy of the British Museum (Natural History).

320 JOHNSONIA, No. 38 Pinnidae

(T. E. Pulley; USNM); San Luis Pass, Galveston (T. E. Pulley); Matagorda Bay
(SDM: CAS: USNM): Indianola (USNM): Aransas Bay (J. L. Baughman; SIO;
USNM);: Port Isabel (E. Weisenhaus; R. Parker); Brownsville (MCZ); 60 miles S of
Brownsville in 12-14 fathoms (C. L. Branch). Cuspa: Varadero Beach, Matanzas (USN M).
Jamaica: Kingston (MCZ; ANSP; USNM); Rock Fort and Port Royal (USNM).
HisraNniota: Port au Prince, Dépt. de Quest; Les Cayes and Torbeck, Dépt. du Sud,
all Haiti (all USNM). Puerro Rico: Bahia de Afiasco (MCZ); Punta Algorrobo and
Punta Guanajibo (both G. Warmke); Mayagiiez (MCZ; G.Warmke). Viren Is_Lanps:
St. Thomas (MCZ: ANSP: USNM; SDM): Gorda Sound, Virgin Gorda (R. Foster).
Lesser ANTILLES: off Fort George, Scarborough, Tobago in 36 fathoms (R. Foster);
Pointe & Pierre; La Brea; and Erin, all Trinidad (all MCZ); Port of Spain, Trinidad
(ANSP). Carisppean IsLtanps: Curacao (USNM). Mexico: Tuxpan, Veracruz (T.E.
Pulley); Veracruz, Veracruz (ANSP). CoLompia: Cartagena (CAS; USNM); Puerto
Colombia (USNM). VeENeEzvuELA: Barcelona (USNM): Campano, Estado Suerre
(ANSP). Brasit: Rio de Janeiro(MCZ; ANSP): Praia Grande, Itanhaem, Estado de
Sio Paulo (M. J. Oliveira); Peruhybe, Itanhaem, Estado de Sao Paulo (H. S. Lopes).
Urvucuay: Cape Polonia, Dept. Rocha(USNM). ArGENTINA: Rio Negro, Patagonia
(USNM).

Atrina (Servatrina) serrata Sowerby
Plates 170; 171

Pinna serrata Solander 1786, Catalogue of the Portland Museum, pp. 71, 165 [nomen nudum].

Pinna serrata Sowerby Feb. 1825, Catalogue of Shells of Earl of Tankerville, London, p. 23, appendix, p. v;
“Solander’ Sowerby Nov. 1825, The Genera of Recent and Fossil Shells, Liv. 26, Pinna, (no locality given) ;
Reeve 1841, Conchologia Systematica 1, pl. 103; Reeve 1859, Conchologia Iconica 11, Pinna, pl. 34, fig.
65 (West Indies).

Pinna squamosissima Philippi 1849 [in] Roemer, Texas, Bonn, p. 454 (near Galveston, Texas).

Pinna seminuda “Lamarck” Reeve 1858, Conchologia Iconica 11, Pinna, pl. 2, fig. 2 (South Carolina); non

Pinna seminuda Lamarck 1819.
Pinna muricata ‘Linné’ Holmes 1860, Post Pleiocene Fossils of South Carolina, Charleston, S.C., p. 15,
pl. 3, fig. 3 (Simmons’; Abbapoola [Abbapoola Creek, Johns Id.], both near Charleston, South Carolina);

non Pinna muricata Linné 1758.

Description. Shell reaching 295 mm. (about 114 inches) in length, wedge-shaped, in-
flated, thin, fragile and sculptured with numerous imbricate, radiating ribs. Shell trans-
lucent, ranging in color from a light tan to a medium greenish brown. Surface, between
the rows of spines, somewhat glistening. Sculpture remarkably uniform for this genus.
On the posterior slope it consists of about 30 inconspicuous ribs which are indicated main-
ly by the imbrications arming them. Sculpture on the ventral slope consisting of concen-
tric growth lines and fine imbrications. Posterior margin usually truncate with a distinct
angle at the union of the posterior and ventral slope. Dorsal margin straight to slightly
concave. Ventral margin convex posteriorly and concave anteriorly. Interior of the valves
iridescent, light tan to greenish brown in color. Nacreous layer thin and usually extending
over three fourths the length of the valve. Posterior adductor muscle scar nearly circular
in outline and set well within the nacreous layer. Anterior adductor muscle scar small,
nearly as wide as the anterior end of the valve.

Western Atlantic JOHNSONIA, No. 38 321

length width
295 mm. 167mm. _ Sanibel, Florida
245 136 Shackelford Id., North Carolina
227 112 12 miles S of St. Augustine, Florida
180 86 Port Isabel, Texas

Types. The holotype of Pinna serrata Sowerby is in the British Museum (Natural
History), no. B3054 from the Tankerville Collection. The type locality is here desig-
nated as Charleston, South Carolina, a locality from which a great deal of material was
sent to Europe at that time. Weare grateful to I.C.J.Galbraith of the British Museum
for the figure of the holotype (Plate 170). The location of the type of Pinna squamosis-
sima Philippi is unknown to us. The type of Pinna seminuda ‘Lamarck’ Reeve is in the
British Museum and that of P. muricata Holmes is in the Charleston Museum, Charles-
ton, South Carolina.

Remarks. Atrina serratais close in its relationship to A. seminuda Lamarck; however,
it may be readily distinguished by its thinner shell, more numerous, crowded ribs, and
low imbricate spines. In A. seminuda the spines are usually tubular and slightly recurved.
The posterior adductor muscle scar of seminuda is usually smaller, proportionately, than
that of serrata and the nacreous layer of semimuda extends only about one half to two

Plate 170. Atrina (Servatrina) serrata Sowerby. Holotype of Pinna serrata Sowerby { = Atrina serrata Sowerby r,
A small, fragile specimen showing the row of spines on the dorsal margin typical of young specimens (nat-
ural size). Photograph courtesy of the British Museum (Natural History ).

399 JOHNSONIA, No. 38 Pinnidae

Jor

thirds the length of the valve, while that of serrata extends at least three fourths the
length of the valve. Inaddition, 4. serrata is a far more inflated species: a specimen 240
mm. (about 94 inches) in length had a thickness of 57 mm. (24 inches), compared with a
specimen of A. seminuda of the same length which is only 41 mm. (12 inches) thick when
measuring the distance through the apposed valves in the vicinity of the posterior adduc-
tor muscle.

Range. From Hatteras Inlet, North Carolina, south through the Florida Keys and in
the Gulf of Mexico to Texas and south to Colombia. It is apparently rare in the West
Indies, having been recorded only from Cuba, Puerto Rico and Trinidad. The paucity
of records in Central America is probably due to lack of collecting.

Specimens examined. Norra Carouina: Point Buxton, Cape Hatteras (USNM);
Hatteras Island, Hatteras Inlet (J. K. Howard); Shackleford Island (MCZ); Fort
Macon, Beaufort (USNM); Bogue Island (MCZ); Cape Fear (ANSP). Sours Caro-

Plate 171. Alrina (Servatrina) serrata Sowerby. Fig. 1. From Mitchell’s Beach, Gulfport, Florida. Inner view
of valve to show the extent of the nacreous layer and the position of the muscle sear. Fig. 2. From Sanibel
Island, Florida. External view of the valve showing the numerous fine, radiating ribs (both about $ natu-
ral size).

Western Atlantic JOHNSONIA, No. 38 323

LINA: Myrtle Beach (ANSP); Isle of Palms (E. B. Richardson); Charleston (MCZ;
USNM): Ashe Island, mouth of Edisto River (ANSP). Groreia: Sea Island, Glynn
Co. (MCZ; ANSP); St. Simons Island (ANSP; USNM): South Beach, Brunswick
(ANSP). Frorimpa: Fernandina (ANSP); Amelia Island (MCZ; ANSP; USNM);
Pablo Beach, near St. John River (MWCZ; ANSP); Atlantic Beach, near Jacksonville
(MCZ); South Beach, Anastasia Island, St. Augustine (ANSP); 12 miles S of St.
Augustine (MCZ); 7 miles S of Matanzas River, St. Johns Co. (ANSP); Ormond
Beach: Coronado Beach; New Smyrna; Cape Canaveral: off Beacon D, The Elbow,
Key Largo in 50 and 100 fathoms; 5 miles SE of Sombrero Light, Marathon, Key
Vaca (all MCZ); Cape Romano; Fort Meyers Beach (both ANSP); Sanibel Island
(ANSP; MCZ: USNM: SDM: J. Schwengel; G. Kline); Captiva Island (J. Schwen-
gel); Boca Grande, Gasparilla Island (ANSP); Long Key, off Sarasota (MCZ): Anna
Maria Key, Sarasota Bay (USNM); Mullet Key, Tampa Bay (ANSP); Pass-a-Grille
(CAS); Mitchell’s Beach, Gulfport (MCZ); Clearwater Bay (ANSP); Cedar Keys
(SDM); St. Joseph’s Bay, Calhoun Co. (ANSP); Port St. Joe (MCZ; USNM): Pan-
ama City (USNM); Pensacola (SDM). Mississtppr: Horn Island (D. Moore). Loutsi-
ANA: Breton Island (R. Parker); Mississippi Delta (K. Johnstone). TExAs: Galveston
(T. E. Pulley); Matagorda Bay (CAS; USNM); Port Aransas(MCZ; ANSP); Padre
Island, near Port Isabel (R. Parker): Port Isabel (E. Weisenhaus). Mexico: Tuxpan,
Veracruz (T. Pulley). Cotompia: Cartagena (CAS). Purerro Rico: Punta Guanajibo,
4 miles S of Mayagiiez (G. Warmke). Cupa: Varadero Beach, Matanzas (ANSP).
Lesser ANTILLES: off Fort George, Scarborough, Tobago in 36 fathoms (MCZ). Near
Erin River, Trinidad (H. G. Kugler).

324 JOHNSONIA, No. 38 References

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Simmonds, P. L. 1879. The Commercial Products of the Sea. Griffith and Farran, London, pp. 306-310
(viii t484 pp).

$26 JOHNSONIA, No. 38 References

Suto, Ryoichi, von 1938. Zur Frage des Mangans in der Steckmuschel. Ein Beitrag zur Kenntnis des Pinna-
globins. Japanese Journal of Zoology, Transactions and Abstracts 8, No. 1, pp. 123-127.

Thiele, J. 1934. Handbuch der Systematischen Weichtierkunde 2, pp. 803-804. Gustav Fischer, Pub.,

Jena.

Vokes, H. E. 1939. Molluscan Faunas of the Domengine and Arroyo Hondo Formations of the California
Eocene. Annals New York Academy Sciences 38, p. 50.

Wilkins, G. L. 1953. Notes from the British Museum, |. Pimna. Proceedings Malacological Society London
30; pp. 25—29.

Winckworth, R. 1929. Marine Mollusea from South India and Ceylon. III: Pinna. With an index to the
Recent species. Proceedings Malacological Society London 18, pp. 276-297.

—— 1936. Marine Mollusca from South India and Ceylon. X: Further notes on Pinna. Proceedings Mala-
ecological Society London 22, pp. 20-23.

Yates, J. 1843. Textrinum Antiquorum. Taylor and Walton, London, pp. 152-159.

Yonge, C. M. 1953. Form and Habit in Pinna carnea Gmelin. Philosophical Transactions of the Royal So-
ciety London. Series B., Biological Sciences, no. 648, 237, pp. 335-374.

Zittel, Karl A. von 1900. Textbook of Palaeontology 1, pp. 368-369, fig. 621. MacMillan & Co., Ltd., Pub.,

London.

The Works of Georgius Everhardus Rumphius
1626-1693

The work of Rumphius though published in 1705, some fifty-three years prior to the
appearance of the tenth edition of Linné’s Systema Naturae, is of great importance to
all malacologists because of the many references which Linné made to it. At the time
Linné wrote the tenth edition of the Systema Naturae in 1758, the starting point of our
binomial system of nomenclature, he did not have in his collection a number of the spe-
cies which he named in his book. In addition, the Systema Naturae was not illustrated
and consequently, to aid in the understanding of his species, Linné referred to one or
several illustrations in other works. One of those most quoted by Linné was that of
Rumphius, D’Amboinsche Rariteitkamer, referred to by Linné as ‘‘Rumph. mus.”’
Consequently, in many cases the figures in Rumphius have been selected by subsequent
authors as the ‘‘type figures’* of Linné’s species.

Rumphius, or Georg Eberhard Rumpf_ as his name was known in Dutch, was born
in Hanau, a small town in Hesse, Germany in 1626 and apparently moved to Holland.
According to Bickmore, Rumphius ‘‘having studied medicine, at the age of twenty-
eight went to Bavaria, entered the merchantile service of the Dutch East India Company,

Western Atlantic JOHNSONIA, No. 38 327

and thence proceeded to Amboina, where he passed the remainder of his life. At the age
of forty-two, while contemplating a voyage back to his native land, he suddenly became
blind and therefore never left his adopted island home; yet he continued to prosecute
his favorite studies in natural history till his death, which occurred in 1693, when he had
attained the ripe age of sixty-seven.’’ According to Swainson, he was at one time gov-
ernor of Amboina.

The first edition of his great work on the shells of Amboina ‘‘D’ Amboinsche Rariteit-
kamer’’ or Amboina Curiosities was published in folio edition at Amsterdam in 1705,
twelve years after his death, and for many years it was the standard work on the mol-
lusks of this area. Another edition, also in Dutch appeared in 1741 using the same plates.
It is this edition that is most commonly seen today. This edition is divided into three
parts or books, the first including pages 1-54 and plates 1-16 covered the crustaceans
and echinoderms, the second including pages 55-194 and plates 17-49 dealt with the
marine mollusks of Amboina and the third, covering pages 195-840 and plates 50-60
described the minerals, stones and artifacts. The illustrations are for the most part good
for the time though many of the shells are reversed as some of the illustrations were ap-
parently cut directly on wood blocks and not from a mirror image, a practice which later
became universal with all engravers. ‘To make such reversed specimens look natural one
should look at them in a mirror.

A German edition of this work entitled “‘Amboin Raitiéiten-Kammer’’ which was
translated by Miiller and emended by Chemnitz was published in Wien in 1766. In this
edition a number of the plates and much of the text were changed. It cannot, therefore,
be used in conjunction with Linné’s work.

The ‘‘Thesaurus Imaginum Piscium Testaceorum, ete.’’ by Rumphius was published
in 1711. This book was a folio edition containing 60 plates, the same used in the “‘D’
Amboinsche Rariteitkamer’’ but the text was reduced to the plate captions and an index.

According to Bickmore, when Holland became a province of France in 1811, attempts
were made to make Paris the center of science and literature in Europe and the Rumphius
collection was taken from Leyden to Paris. ‘Though it was later returned, much damage
had been done to the collection during the two transfers and a large portion of the speci-
mens had disappeared. Louis Agassiz apparently knew of this loss and realized that an
understanding of many of Linné’s species depended upon a knowledge of the specimens
which Rumphius had figured in his great work. Consequently, in 1865 he sent Albert S.
Bickmore, one of his students, to Amboina for the express purpose of duplicating the
Rumphius Collection. Much of this material is now in the Museum of Comparative
ZoGdlogy.

Rumphius also wrote extensively on the botany of Amboina and he is generally cred-
ited with having laid the foundations of botanical and zoological research in the East
Indies. During his trip to Amboina, Bickmore located the grave of Rumphius which
was marked by a small square pillar erected in 1824 by G. A. Phillipus, the royal gover-
nor to replace the original marker which had been destroyed.

Bickmore, Albert S. 1869. Travels in the East Indian Archipelago. D. Appleton and Company, New York,
553 pages, 36 plates.
—Rutu D. Turner

328 JOHNSONIA, No. 38 Book Review

Book Review

Olsson, A. A., A. Harbison, W. G. Fargo and H. A. Pilsbry 1953. Pliocene Mol-
lusca of Southern Florida with special reference to those from North Saint Petersburg.
The Academy of Natural Sciences Philadelphia: Monographs—Number 8, pp. viii+1-
457: 65 plates; 2 text figs.; 2 maps. From the time of its discovery in 1988 by A. P.
Cales, the Pliocene fossil bed of North Saint Petersburg has been well explored and col-
lected. The site represents the farthest known northwest extension of the Caloosahatchee
formation of southern Florida which is well known as a result of the work of Dall in
1890 to 1903. This recent publication is a study of the mollusks, predominantly marine,
which have been collected at the St. Petersburg site. Several authors have reported on
the material: Part I, by Olsson and Harbison, is a systematic consideration of the Classes
Palecypoda, Scaphopoda and Gastropoda; Part II, on the gastropod family Turridae,
was done by Fargo who also gave a summary of the geology of the Saint Petersburg re-
gion which is included in the general introduction; Parts III-A and III-B, by H. A
Pilsbry are devoted, respectively, to studies of the gastropod family Vitrinellidae, and
the freshwater gastropods and pelecypods.

As noted by Fargo in the introduction to the study, the Pliocene shells of Saint
Petersburg were readily distinguished from Recent or Pleistocene specimens in the area
because of their distinctive buff or pale reddish color. For this reason placement in geolo-
gic time was more certain than it may be in other areas. Mollusks of the Pliocene which
also extend into the Recent were found to comprise 32.49% of the species studied from
this site. It was pointed out in the introduction that one difficulty in a study of this
type is the determination of the actual number of fossil species which extend into the
Recent. This is particularly difficult because of the lack of a complete census of the liv-
ing mollusk fauna of the West Indian and Gulf areas.

The authors have named 10 new genera, 30 subgenera and 1 new section of mollusks.
Of the 517 species and subspecies treated in this report, 1837 were described as new. This
systematic study will be found extremely useful by malacologists interested in either
fossil or recent mollusks of the Florida area.

The Pyramidellidae from the Saint Petersburg collection were sent to the U.S. National
Museum and were treated by Dr. Paul Bartsch in his 1955 publication: ‘‘The Pyrami-
dellid Mollusks of the Pliocene Deposits of North St. Petersburg, Florida’’; Smithson-
ian Miscellaneous Collections 125, no. 2, pp. 1-102, 18 plates.

—JosEPH ROSEWATER

oe em" MUS. CSAP. Z68L

LIBRARY
JUN 25 1959

JOHNSONIA uNIVERSIT

Published by
THe DEPARTMENT oF MOLLUSKS
Museum of Comparative Zodlogy, Harvard University
Cambridge, Massachusetts

JUNE 26, 1959 REVIEW NUMBER VOL. 3, NO. 39

THE GENUS CONUS IN THE WESTERN ATLANTIC
BY
WILLIAM J. CLENCH

Conus fosteri Clench and Aguayo

Conus fosteri Clench and Aguayo 1942, Johnsonia 1, no. 6, p. 34, plate 12, fig. 5 (off Sagua la Grande,
Santa Clara [Las Villas] Cuba).

Specimens examined. VENEZUELA: Atlantis, cruise 240, off Cabo Codera (N. Lat. 11°;
W. Long. 66°1’) in 150 fathoms (W. Athearn).

This extends considerably the range of this species. Heretofore it was known only
from Cuba.

THE GENUS SCONSIA IN THE WESTERN ATLANTIC
BY
WiLuiaM J. CLENCH

Sconsia striata Lamarck
Plate 172

Sconsia striata Lamarck. Clench and Abbott 1943, Johnsonia 1, no. 9, p. 6.

Plate 172. ‘Sooasin barbudensis Higgins and Marrat (=Sconsia striata Lamarck). Holotype, 15 fathoms, off
Barbuda, Lesser Antilles (slightly enlarged).
329

330 JOHNSONIA, No. 39 Columbarium

At the time of our original study of this genus, it was impossible to get photographs
from Europe. Through the kindness of Mr. W. K. Ford, Keeper of Invertebrate Zool-
ogy of the City of Liverpool Public Museums we have obtained a photograph of the
holotype of Sconsia barbudensis Higgins and Marrat. This species is a synonym of
Sconsia striata Lamarck. The measurements of the holotype are: length 67 mm., width
38 mm.

THE GENUS COLUMBARIUM IN THE WESTERN ATLANTIC
BY
WILLIAM J. CLENCH

Through the kindness of Mrs. Roy C. Athearn I am privileged to describe the follow-
ing species. The specimens were obtained by her son, William Athearn during a cruise
of the Atlantis off the coast of Venezuela.

Columbarium brayi, new species
Plate 173, figs. 1-2

Description. Shell medium in size, reaching 51 mm. (2 inches) in length, imperforate,
rather thin, dull and carinated. Color a dull light yellowish brown. Whorls 9 and sharply
carinated. Spire somewhat depressed. Suture deeply impressed. Aperture subquadrate.
Outer lip simple. Parietal wall thinly glazed. Columella long and straight, extending to
the end of the long siphonal canal. Sculpture consists of numerous fine, spiral cords
which are interrupted by the growth lines. Embryonic whorls two, smooth and glass-
like. Periostracum thin with short axial blades. Operculum unknown.

length width
51.0 20mm. _ Holotype
Bilao 18 Paratype

Types. Holotype, Museum of Comparative Zoology, no. 221601, from Atlantis, cruise
240, off Cabo Codera, Venezuela (N. Lat. 11°; W. Long. 66°1’) in 150 fathoms, W.
Athearn collector, Nov. 1, 1957. A single paratype from the same station.

Remarks. This species does not appear to be closely related to any other species of
Columbarium in the Western Atlantic. Its larger size and lack of spines differentiate it
from C. atlantis Clench and Aguayo, its larger size and depressed spire distinguish it from
C. bermudezi Clench and Aguayo and its size and smooth carina from C. sarissophorum
Watson. This last species has a uniformly serrated carina (see Johnsonia 1944, 1, no. 15,
Peo)

This species is named for Captain W. Scott Bray of the Atlantis.

Columbarium atlantis Clench and Aguayo
Columbarium atlantis Clench and Aguayo 1944, Johnsonia 1, no. 15, p. 2, pl. 1, fig. 4 off Matanzas, Cuba).

Specimens examined. Cusa: Atlantis, station 3484, off Sagua la Grande, Las Villas
(N. Lat. 23°10’; W. Long. 79°35’) in 260 fathoms.

Western Atlantic JOHNSONIA, No. 39

Plate 173. Columbarium brayi Clench. Fig. 1. Holotype. Fig. 2. Paratype. 150 fathoms off Cabo Codera,
Venezuela (2.8x).

THE GENUS MUREX IN THE WESTERN ATLANTIC
BY
WILLIAM J. CLENCH

Murex (Murex) finlayi Clench
Plate 174, figs. 1-3

Murex (Murex) finlayi Clench 1955, Breviora, Museum of Comparative Zoology, no. 44, pp. 1-3, text figs.
1-3 (Matanzas Bay, Cuba).

Description. Shell reaching 98 mm. (3% inches) in length, rather solid in structure and
moderately spinose. Whorls 10 and moderately convex. Nuclear whorls glass-like and
smooth. Color brownish yellow to brownish cream with a few spiral threads of a darker

332 JOHNSONIA, No. 39 Murex

brown which follow the crests of the cords. Spire moderately extended. Suture irregular
and deeply indented. Aperture subcircular and slightly oblique. Parietal lip glazed, ad-
herent to the body whorl and fairly thick. Palatal lip crenulated and slightly thickened.
Siphonal canal greatly extended, sometimes nearly half the length of the shell, usually
curved upward and to the left when viewed dorsally. Two or three previously formed
stages of the siphonal canal remain as scale-like spines. The sculpture consists of three
equidistant varices, each supporting a rather large shoulder spine and maybe two or three
smaller spines between the shoulder and the base of the shell. The varices on each whorl
are more or less aligned with the varices on the whorl above. Between the varices there
are two to four axial ridges which are strongly nodulose, Spiral sculpture consisting of
numerous thread-like cords. The varices are formed by two arched plates, the plates on
the aperture side being somewhat smaller, the outer (first formed) leaving a flange or
web-like ridge as a crest on the varix. This is usually broken away on the early varices
or remains as a series of small blade-like spines.

length width whorls
87 30 10 Holotype
93 31 10 Paratype
85 33 10 Paratype
82.5 34 10 Paratype

Types. Holotype, Museum of Comparative Zoology, no. 189939 from shell trap,
Matanzas Bay, Cuba, in 100 fathoms. Paratypes from the same locality in the Museum
of Comparative Zoology and the collections of C. J. Finlay and E. H. Monroe.

Plate 174. Murer (Murex) finlayi Clench. Figs. 1 and 3. Paratypes. Fig. 2. Holotype. 100 fathoms,
Matanzas Bay, Cuba (all slightly enlarged).

Western Atlantic JOHNSONIA, No. 39 333

Remarks. 'This species appears to be rather closely related to both Murex antillarwm
Hinds and Murex beaui Fischer and Bernardi. From M. antillarum it differs by having
fewer and much larger axial costae, having fewer spines, and in possessing a webbing
along the varices and greatly extended siphonal canal. From M. beauii it differs by hav-
ing fewer and much larger axial costae, shorter spines, and not having the extensive web-
bing which characterizes M. beauw. This present species may also be related to M.
aguayot Clench and Farfante but it differs from aguayoi by being larger, having the dif-
fused brownish yellow coloration, fewer spines, larger axial costae and by having the
webbing on the varices.

The development of the webbing and the greatly extended siphonal canal probably
allows these mollusks to exist on a rather soft muddy bottom.

The subgenus Murevx in which this species is included is a very complex group and of
very wide distribution in nearly all tropical and south temperate seas.

Murex (Murex) pulcher 4. Adams

Murex (Murex) pulcher A. Adams. Clench and Farfante 1945, Johnsonia 1, no. 17, p. 23, pl. 12, figs. 1-4.

Specimens examined. Brasiu: Hassler Voyage off Salvador (Bahia) in 40 fathoms
(S. Lat. 11°49’; W. Long. 37°20’) (MCZ).

This record extends the range of this rare species from the Lesser Antilles to the coast
of north central Brasil.

Murex (Phyllonotus) pomum Gmelin

Murex (Phyllonotus) pomum Gmelin. Clench and Farfante 1945, Johnsonia 1, no. 17, p. 26, pl. 14, figs. 1-3.

Specimens examined. Fioripa: St. Andrews Bay, Panama City (R. Work). Texas:
5 miles W of Rockport (MCZ), Port Aransas (T. Pulley).

Murex (Phyllonotus) pomum margaritensis Ablott
Plate: 175, figs. 1-2

Murex imperialis Swainson 1831, Zoological Illustrations (2) 2, pl. 67 (Island of Margarita [Venezuela] ).
Non Murex imperialis Fischer 1807.

Murex (Phyllonotus) pomum Gmelin. Clench and Farfante 1945 [in part] Johnsonia 1, no. 17, p. 26.

Murex margaritensis Abbott 1958, Acad. Nat. Sci., Philadelphia, Monograph no. 11, p. 61, pl. 1, figs. n
and o. [New name for Murex imperialis Swainson, non Fischer].

In our Johnsonia number on Murex (1945, 1, no. 17, p. 26) we considered Murex
imperialis to be a synonym of Murex pomum Gmelin. At that time we had seen no
specimens so our judgment was based only upon descriptions and figures in Swainson,
Reeve and Kiener, all of whom failed to show the deep pink coloration of the aperture.
Abbott has mentioned several characters which distinguish this form from pomum. When
a series of this form is examined, all of these characters, other than the pink coloration,
are found in both pomum and imperialis. 'The number of varices on the body whorl vary
in both forms, but the variation appears to be similar in both pomum and imperialis.
There are two rows of spines on most specimens of imperialis we have seen, not one row
as stated by Abbott.

334 JOHNSONIA, No. 39 Murex

Many specimens of ¢mperialis lack the brown patch on the parietal area, but in others
it is present. Most specimens of imperialis are less attenuated than pomum, but even this
character is occasionally duplicated in large specimens of pomum.

length width
96 mm. 70 mm. Margarita Id.
68 43.5 Erin River, Trinidad

Range. Margarita Island, Venezuela east to Trinidad.

Specimens examined. VENEZUELA: Margarita Island (MCZ). Lesser ANTILLES: near
Erin River; Guayaguagare Beach; Magueripe Bay; Couva Bank, Gulf of Paria, all

Trinidad (all H. G. Kugler).

Plate 175. Murex pomum margaritensis Abbott. Fig. 1. With a clear pink aperture. Fig. 2. With the brown
parietal spot. Margarita Island, Venezuela (about natural size).

THE GENERA HEMITOMA AND DIODORA
IN THE WESTERN ATLANTIC
BY
Rurn D. Turner

Hemitoma is a small and relatively little known genus in the Fissurellidae. It is close
to Hmarginula, but in Hemitoma the anal notch is reduced or lacking but there is an
internal anal groove. A key to the genera and subgenera of the Fissurellidae of the

Western Atlantic appeared in Johnsonia 2, no. 24, p. 94.

Western Atlantic JOHNSONIA, No. 39 335

Genus Hemitoma Swainson

Hemitoma Swainson 1840, Treatise on Malacology, p. 356; Iredale 1915, Transactions New Zealand In-
stitute 47, p. 433.

Subemarginula Gray 1847, Proceedings Zoological Society London 15, p. 147 (type species, Patella octo-
radiata Gmelin, by monotypy).

Siphonella Issel 1869, Malacologia del Mar Rosso, p. 232 (type species, Emarginula (Siphonella) arconatii
Issel, by monotypy).

Hemitoma ‘Rafinesque’ Scudder 1882, Nomenclator Zoologicus, Part I, Supplemental List. Bulletin United
States National Museum, no. 19, p. 159 [error for Hemiloma Rafinesque |.!

Type species, H. [Hmarginula] tricostata Sowerby [=octoradiata Gmelin], by mono-
typy.

Shell patelliform, small to medium in size, solid in structure, elliptical to broadly oval
in outline, depressed to highly conic. Sculpture consisting of radial ribs and in some spe-
cies both radial ribs and concentric ridges. Anterior rib usually the most prominent and
often with a small notch at the ventral margin. Muscle scar horseshoe shaped with the
anterior ends recurved toward the apex as triangular spurs. The scar left by the attached
area of the mantle anterior to and between the spurs is probably responsible for the de-
scription of the muscle scar in this group as being trilobed. Radula with a moderately
wide central tooth which has a single cusp without denticles. The first three lateral teeth
are similar but much narrower. The fourth lateral is large, nearly covering the third lat-
eral when in its normal position. At the base of this tooth and between it and the first
marginal tooth there is a small squarish plate or tooth which is quite flat except for a
central thickened ridge. This plate was first figured by Odhner (1982, fig. 41). This is
similar to what Robertson (1958) has shown in the radula of 7’icolia and referred to as
a latero-marginal plate. It is interesting that this structure should appear in two such
unrelated groups, or has it been overlooked in other families? The marginal teeth of
Hemitoma range in number from 30 to over 60, they are long, narrow, with a single
cusp and numerous denticles.

Gray 1847 gave credit to de Blainville for the name Subemarginula with Patella
octoradiata Gmelin as the type. However, de Blainville used this name only in the ver-
nacular—‘Les subemarginules.’” Though Pilsbry (1891) used Subemarginula de Blain-
ville 1825 with Hmarginula emarginata de Blainville as the type species, the first accept-
able use of the name is that of Gray 1847. Hemitoma Swainson 1840, having seven
years priority, is the name generally accepted for this genus today. Iredale (1915, p. 433)
has given a detailed discussion of this problem. However, he was in error in using Sub-
emarginula Gray for octoradiata alone, and considering Hemitoma a distinct genus with
tricostata Sowerby as type. This species is a synonym of octoradiata and Subemarginula
an objective synonym of Hemitoma.

The oldest known American species in this genus is Hemitoma (Hemitoma) sciera which
was described by Woodring (1928, p. 475) from the Miocene of Bowden, Jamaica. This

‘In the Preface to his book Scudder stated: “‘Where the entry has been furnished by an author or other
correspondent his name is appended in italics as its sponsor; the name Agassiz is appended to all entries
copied from his manuscript additions to the Nomenclator of 1846.°* Such was the case with Hemiloma Rafines-
que. However, when checking Agassiz’ Nomenclator, we find that he had not included Hemiloma Rafinesque
and it was this to which he was referring in his manuscript and Scudder made the error in copying it.

336 JOHNSONIA, No. 39 Hemitoma

species is very close to the recent species H. octoradiata Gmelin which is common
throughout the West Indies. The only fossil records from the mainland are those of
Hemitoma (Montfortia) retiporosa Dall (=emarginata de Blainville) from the Pliocene
of Florida. Wenz (1988) records the genus as first appearing in the Eocene.

Odhner (1932) has discussed the anatomy of Hemitoma in relation to the other genera
in the Fissurellidae and has illustrated the anatomy of H. octoradiata.

This genus is world wide in distribution in warm temperate and tropical seas.

Subgenus Hemitoma Swainson

Species in this subgenus are characterized by having the sculpture predominantly
radial, with strong primary and secondary radial ribs. The concentric sculpture is weak,
usually consisting only of growth lines. The anal notch is lacking except in some very
young depressed forms but the internal anal groove is well marked. The apex is nearly
central, usually blunt and inclined only slightly posteriorly.

There is only a single recent species in this subgenus in the Western Atlantic.

Hemitoma (Hemitoma) octoradiata Gmelin
Plates 176;.177

Patella octoradiata Gmelin 1791, Systema Naturae, ed. 13, p. 3699 (Insulas Americae mediae).

Emarginula tricostata Sowerby 1824, The Genera of Recent and Fossil Shells, no. 21, Emarginula, fig. 6
(no locality given).

Emarginula listeri Anton 1839, Verzeichniss der Conchylien, p. 27 (refers to Lister, pl. 532, fig. 11—Bar-

bados).

Emarginula clausa d’Orbigny 1842 [in] Sagra, Histoire d’Ile de Cuba, Mollusques 2, p. 194, pl. 24, figs.
34-36 (Cuba).

Subemarginula octoradiata Gmelin. Gray 1847, Proceedings Zoological Society London 15, p. 147; A. Adams
1851 [1852], Proceedings Zoological Society London, p. 90; Pilsbry 1891, Manual of Conchology 12, p. 273.

Emarginula depressa Sowerby 1868, Thesaurus Conchyliorum 3, p. 219, pl. 247, figs. 64, 65,68 (no locality
given).

Emarginula guadaloupensis Sowerby 1863, Thesaurus Conchyliorum 3, p. 219, pl. 247, fig. 69 (Guadeloupe
Island, on basis of name).

Hemitoma octoradiata Gmelin. Woodring 1928, Carnegie Institution of Washington Publication, no. 385,

p. 457.
Hemitoma rubida A. H. Verrill 1950, Nautilus 63, p. 126, pl. 9, figs. 2, 2a (Canefield Point, Dominica,
B.W.I.).

Description. Shell solid, reaching 80 mm. (14 inches) in length, patelliform and radially
sculptured. Extremely variable in shape, ranging from nearly circular to elliptical in
outline, the height varying from about { to nearly the length of the shell. Apex sub-
central and pointed slightly posteriorly. Anterior slope slightly to strongly convex.
Posterior slope slightly concave immediately below the apex, then varying from nearly
straight to rather strongly convex. Radial sculpture consisting of eight unbranched, ir-
regularly nodulose primary ribs which extend from the apex to the basal margin. In
young specimens the primary ribs are very prominent and project beyond the margin
producing a stellate appearance. As the shell grows, secondary and often tertiary ribs
are produced between the primary ribs, and the margin becomes finely crenulate. Con-
centric sculpture consisting of irregular growth lines only. Color a light tan or gray to
greenish brown. Periostracum thin, medium golden brown in color and usually seen only

Western Atlantic JOHNSONIA, No. 39 337

on young specimens. Interior of shell a glossy olivaceous green to purplish brown with
a white margin and a white area at the apex. Anal groove narrow and deep, extending
inward about $ to % the length of the anterior slope.

The soft parts of this species are extraordinarily beautiful. The main portion of the
mantle and foot are a medium blue-green, the base of the foot being somewhat lighter.
This coloration increases to an intense turquoise near the edge of the mantle, while the
edge proper is a vivid magenta. The base of the tentacles and the head are also turquoise
while the ends of the tentacles and a circle around the end of the proboscis are magenta.

The radula is similar to that of emarginata though the central tooth is somewhat
broader and there are far more marginal teeth, there being from over 45 to 60 marginals
depending upon the age and size of the specimen.

length width height

18.5 mm. 12mm. 7.5mm. Arthurs Town, Cat Island, Bahama Ids.
23.5 20 20 a ie, et Ord

19 15 13 Whitehouse, Jamaica

23 23 15.5 >i Py

22 18 8 Barbados

30.5 24.5 15.2 -

24.5 20 9 Clifton Harbour, Union Island, Grenadines
29.5 25 the Tortugas, Florida

Types. Patella octoradiata Gmelin as originally described was a composite species.
Gmelin’s description applies quite well to this species, but the references are certainly
composite. His first reference was to Lister, plate 532, fig. 11 and this was used by A.
Adams in 1852 to indicate Gmelin’s species. Pilsbry followed Adams and selected

pon meenenerereenee enema

3

Plate 176. Hemitoma octoradiata Gmelin. Series of specimens selected to show variation in shape and sculp-
ture. Fig. 1. Marina Cay, south of Great Camanoe Island, Virgin Islands. Fig. 2. Cayo Francés, Cai-
barién, Cuba. Fig. 3. Clifton Harbour, Union Island, Grenadines. Figs. 4-7. Cienfuegos Harbor, Cuba.
Fig. 8. Dundas Town, Great Abaco, Bahama Islands. Fig. 9. Clifton Harbour, Union Island, Grenadines.
Fig. 10. Santa Barbara de Samana, Dominican Republic. Fig. 11. Internal view, Marina Cay, south of
Camanoe Island, Virgin Islands.

Figs. 1-3. Side view to show variation in height (anterior to the left). Figs. 4-10. Dorsal view to show
growth series (anterior end downward). Fig. 10. A specimen with numerous radial ribs (all 12x).

338 JOHNSONIA, No. 39 Hemitoma

Lister’s figure as the type. As Gmelin gave a very indefinite locality for this species, the
type locality is here restricted to Barbados, the locality given on Lister’s figure. The
type of Hmarginula clausa d’Orbigny is in the British Museum according to Gray
(1854). The types of H. guadaloupensis Sowerby and E. depressa Sowerby are also
probably in the British Museum (Natural History).

Remarks. This is an extremely variable species as indicated in the description. Speci-
mens may range from those which are quite flat and nearly circular in outline with strong
ribs and deeply crenulated margins to those which are very high and conical with weak
ribs and nearly smooth margins, and there may be any combination of these characters.
As shown in the measurements, the length of mature specimens may Vary in relation to
the height from 1.15 to 2.75 and the width may vary from less than 3% the length to
equal the length. Hemitoma rubida Verrill is one of the flattened, broad and strongly
sculptured types. The anterior rib is usually the most prominent, but this is not always
so and the anal groove which follows it on the internal surface of the shell usually does
not end in a notch at the margin though in occasional young specimens there may be a
very small one. Young specimens are usually somewhat rectangular in outline, depressed
and stellate. Specimens of this species are usually moderately to heavily encrusted with

Plate 177. Radulae of Hemitoma. Fig. 1. Hemitoma emarginata de Blainville. Fig. 2. Hemitoma octoradiata
Gmelin. Fig. 3. Embryonic whorls of H. emarginata de Blainville.

Western Atlantic JOHNSONIA, No. 39 339

coralline algae and other growths and it is often necessary to clean them in order to study
the sculpture. However, the muscle scar, the anal groove and the color pattern on the
inner surface are usually sufficient for identification.

Robert Robertson who has spent a great deal of time collecting and studying in the
Bahama Islands writes as follows concerning this species. ‘‘In the Bahama Islands HZ.
octoradiata is abundant in the vicinity of Elbow Cay, Great Abaco (the windward edge
of the Little Bahama Bank). It usually lives a few inches below low tide mark on ex-
posed rocky shores. The irregular surface of the rock is covered with lithothamnia (chiefly
Porolithon) and in this these limpets form slight depressions. They appear to be seden-
tary. They may also be found under slabs of dead coral (Acropora) in the reef northeast
of Elbow Cay. Occasionally specimens were found under rocks and live corals in shallow
water in sheltered areas. At Bimini, on the leeward side of the Great Bahama Bank,
this limpet is scarce. The rocky shores here are not as exposed as many of those at
Abaco. Feces of this Hemitoma studied at Bimini contained chiefly green algae but also
a copepod and series of marginal teeth from its own radula.”’

Range. From Biscayne Bay, Florida south through the Florida Keys, Bermuda
(Peile 1926), the Bahama Islands, throughout the West Indies and along the north
coast of South America south to Porto Seguro, Brasil.

Specimens examined. Fioripa: Biscayne Bay (MCZ); Pelican Shoals off Boca Chica
Key (J. Schwengel); Middle Sambo Shoals, near Key West (T. McGinty); Sand Key,
Key West; Dry Tortugas (both MCZ). Banama Istanps: West End and Eight Mile
Rock, Grand Bahama (both MCZ); Cooper Jacks Cays, S of Elbow Cay and Dundas
Town, Great Abaco; North Rabbit Cay, Bimini Islands (all R. Robertson); Brown’s
Point, New Providence; Governors Harbour, Eleuthera; Cape St. Maria; Clarence
Town and Simms, all Long Island; Little San Salvador and Russell Creek, Cat Island:
Rum Cay; Matthew Town, Great Inagua (all MCZ). Cuna: Vedado and Playa de
Jibacoa, Habana; Cayo Cruz del Padre and Peninsula de Hicacos, Matanzas (all MCZ);
Cayo Francés, Caibarién, Las Villas; Santa Maria Key, off Punta Alegre, Camagiiey
(both R. Humes); Vita; Guarda la Vaca, Banes; Bahia de Banes; Blue Beach, Guan-
tinamo Naval Base, all Oriente; Punta de los Colorados, Cienfuegos, Las Villas (all
MCZ). Jamaica: Runaway Bay, St. Ann’s; Montego Bay (both MCZ); Whitehouse
(J. K. Howard). Hispanioua: Gonave Island, Haiti (W. J. Eyerdam); Monte Cristi;
Puerto Plata; Puerto Sostia; Santa Barbara de Samana, all Dominican Republic (all
MCZ). Puerro Rico: Mata de la Gata, off La Paguera; E of Boca de Congrejos (both
MCZ). Virecin Istanps: Virgin Gorda: Marina Cay and Guana Island, ‘Tortola (all
M. W. Dewey); St. John; Great St. James Island, St. Thomas (both MCZ); Ham
Bay, St. Croix (M. K. Jacobson). Lesser ANTILLES: Antigua; Guadeloupe; Marigot,
St. Lucia (all MCZ); Barbados (P. G. Kellett); Carriacou Island, Grenadines: Grand
Anse, Grenada (both H. G. Kugler): Buccoo Reef, Tobago (MCZ); Toco, north coast
of Trinidad (H. G. Kugler). Cartmpean IsLanps: Southwest point; Grand Cayman,
Cayman Islands(MCZ). British Guiana: Mouth of Corentyne River (H. G. Kugler).
BrasiL: Porto Seguro (MCZ).

340 JOHNSONIA, No. 39 Hemitoma

Subgenus Montfortia Pécluz

Montfortia Récluz 1843, Revue Zoologique par la Société Cuvierienne 6, p. 259 and 376.

Type species, Hmarginula australis Quoy and Gaimard, subsequent designation, Ire-
dale 1915.

Species in this subgenus are characterized by having a pronounced radial and concen-
tric sculpture with the three anterior ribs usually strongest and by having a small anal
notch at the margin of the anterior rib in addition to the internal anal groove. The apex
is prominent, subcentral and recurved posteriorly.

Hemitoma emarginata de Blainville
Plates 177, 178

Emarginula emarginata de Blainville 1825, Manuel de Malacologie, p. 501, pl. 68 bis, fig. 3 (no locality
given).

Emarginula octoradiata ‘Gmelin’ Sowerby 1863, Thesaurus Conchyliorum 3, p. 219, pl. 247, fig. 66 (no
locality given); non octoradiata Gmelin 1791.

Subemarginula retiporosa Dall 1903, Transactions Wagner Free Institute Philadelphia 3, pt. 6, pl. 55, fig.
3; pl. 60, fig. 17 (Pliocene marl of Shell Creek, Florida). Figured but not described.

Hemitoma retiporosa Dall. Olsson and Harbison 1953, Academy Natural Sciences, Philadelphia, Monographs,
no. 8, p. 360, pl. 48 [not 18], figs. 14, 14a.

Emarginula ostheimerae Abbott 1958, Academy Natural Sciences, Philadelphia, Monographs, no. 11, p. 18,
pl. 1, fig. ce (Old Man Bay, north side of Grand Cayman Island, Cayman Islands).

Description. Shell solid, reaching 25 mm. (1 inch) in length, patelliform and strongly
sculptured. Extremely variable in shape, the basal outline ranging from more or less
quadrate to broadly oval, circular or elliptical. Height ranging from about 1/4 to over
3/5 the length of the shell. Apex subcentral, recurved posteriorly and slightly to the
left. Anterior slope convex. Posterior slope concave just below the apex, then continu-
ing nearly straight or slightly convex. Sculpture consisting of radial ribs and concentric
ridges with nodules formed where the radial ribs and concentric ridges cross, producing
a strong reticulate sculpture. Primary ribs 8 to 10, the anterior rib always being single,
usually prominent and ending in a small notch at the margin. The remaining primary
ribs may be single or compound, the compound ones made up of from 2 to 5 closely
spaced ribs which give the effect of a single broad rib. The spaces between the primary
ribs have from 1 to 5 low secondary ribs. Concentric sculpture consisting of more or less
evenly spaced ridges, the distance between them increasing slightly as the shell grows.
Margin of mature specimens usually moderately to finely crenulate. Young specimens
have a strongly scalloped margin. Shell white, periostracum thin, pale yellow and usu-
ally remaining only in the pits formed by the reticulated sculpture. Interior of the shell
glossy white, often with yellow patches indicating the pits. Anal groove fairly deep and
narrow near the margin, ending in a small notch. It extends as a very shallow groove
to near the apex. Embryonic whorls one and one half, small, smooth and with a light
brown periostracum.

Western Atlantic JOHNSONIA, No. 39 341

length width height
11 mm. 7.3mm. 3mm. Arenas de la Chorrera, Habana, Cuba
20 15 7 Jamaica
91.5 12 12 Fort Jefferson, Tortugas, Florida
23.5 17 8.2 St. Thomas, Virgin Islands
94.5 18.5 tas Off Fowey Rocks, Florida
25.2 19 S22 Nassau, New Providence, Bahama Islands
25.5 15 12.5 Dry Tortugas, Florida

Types. The location of the type specimen of E’marginula emarginata de Blainville is
unknown.

Plate 178. Hemitoma emarginata de Blainville. A series to show variation in shape and sculpture (anterior
end downward). Fig. 1. Jamaica, asomewhat beachworn specimen. Fig. 2. Arenas de la Chorrera, Habana,

?

Cuba. A young specimen with a distinct anal notch. Fig. 3. Tortugas, Florida, a side view to show average
shape in profile (figs. 1, 2 and 4 are somewhat flatter than this; figs. 7 and 8 are about the same and fig.
5 is much higher). Fig. 4. Fowey Rocks, Florida, from an old wreck. Fig. 5. Dry Tortugas, Florida, a
very high narrow form. Fig. 6. Barbados, internal view to show muscle scar and anal groove. Fig. 7.
St. Thomas, Virgin Islands. Fig. 8. Nassau, New Providence, Bahama Islands, from a pier. (All about 2x.)

The holotype of Subemarginula retiporosa Dall is in the United States National Mu-
seum and that of Hmarginula ostheimerae Abbott is in the Academy of Natural Sciences,
Philadelphia, no. 195983.

Remarks. This is a rather rare species and specimens are seldom taken alive. Conse-
quently we know nothing of its ecology. It apparently lives from just below low tide to
depths of perhaps 100 fathoms. Specimens known to have been collected alive which we
have seen, came from a wharf at Nassau, New Providence, Bahama Islands and from a
wreck on Fowey Rocks, Florida. On the basis of the poorly preserved specimen available

342 JOHNSONIA, No. 39 Hemitoma

for study, the anatomy of emarginata appears to be very close to that of octoradiata and
the radula differs mainly in the width of the central tooth and the fewer marginal teeth.

Emarginula ostheimerae Abbott appears to be a young specimen of this species, the
type specimen being almost identical with figure 2 shown in the growth series on Plate
178. Dall figured but never described this species from the Pliocene mar! of Shell Creek,
Florida under the name of Subemarginula retiporosa, and Olsson and Harbison report it
under Hemitoma retiporosa Dall as fairly common in the Pliocene of St. Petersburg,
Florida.

The range of variation in this species is equally as great as that in octoradiata. Many
of the high specimens are also laterally compressed and, as pointed out by Pilsbry (1891),
the anterior and posterior ends of many specimens are slightly raised so that the speci-
men will rock when placed on a flat surface. However, this certainly does not hold true
for all specimens. The extent of the anal groove also varies considerably and is barely
visible in some specimens.

This species is not closely related to any other found in the Western Atlantic. It dif-
fers from octoradiata in having a pronounced reticulate sculpture and compound ribs.

Range. From Lantana, Florida south through the Florida Keys and probably through-
out the West Indies.

Specimens examined. Fioripa: off Lantana in 10 fathoms; off Hollywood in 35-60
fathoms; off Cape Florida, Biscayne Bay in 50 feet; Fowey Rocks; 54 miles NE of
Beacon D, The Elbow, Key Largo in 66-83 fathoms; 6 miles SE of Sombrero Light,
off Marathon in 66 fathoms (all MCZ); Pelican Shoals, Key West in 45 fathoms (J.
Schwengel); Fort Jefferson, Tortugas (MCZ). Bauama IsLanps: W of Picquet Rocks,
Bimini Islands in 10 fathoms (R. Robertson); Northeast shore of New Providence (D. H.
Brown): Nassau, New Providence; Matthew Town, Great Inagua (both MCZ). Cusa:
near Habana (MCZ). Virein Istanps: St. Thomas (MCZ). Lesser ANTILLES: Bar-
bados (MCZ): Fontenary Beach, Grenada (H. G. Kugler).

Genus Diodora Gray

Diodora Gray. Farfante 1943, Johnsonia 1, no. 11, pp. 1-20, pls. 1-6.

The following description of Diodora fragilis is basically a translation of the work of
I. P. Farfante and D. L. Henriquez which appeared after Johnsonia, no. 11 on Diodora
had been published.

Diodora fragilis Farfante and Henriquez
Plate 179, figs. 1—4

Diodora delicata Farfante and Henriquez 1946, Revista de la Sociedad Malacologica ‘Carlos de la Torre’
4, p. 54 (Arenas de La Chorrera, Habana, Cuba).

Diodora fragilis Farfante and Henriquez 1947, ibid. 5, p. 52, figs. 1-4 (new name for D. delicata Farfante
and Henriquez, non E. A. Smith 1889).

Description. Shell small, reaching 10 mm. in length, delicate in structure, depressed
conic, and with the anterior slope considerably shorter than the posterior slope. Base
oval. Anterior slope straight for its entire length, the posterior slope concave just below

Western Atlantic JOHNSONIA, No. 39 343

the apex, the remainder straight to convex. Apex anterior of the center and pierced by
the anteriorly directed orifice. Orifice small, 1 mm. in length and oval. Radial sculp-
ture consisting of numerous rather large, rounded ribs which alternate with very fine
ones. Concentric sculpture consisting of numerous lamina which cross the radial ribs
forming small scales. Margin denticulate. Color grayish white. Interior of shell gray,
glossy and with a series of grooves which correspond to the radial ribs. The callus around
the orifice gray, sharply truncated and excavated posteriorly.

length width height
20mm. 11 mm. 5mm. Holotype

Types. The holotype is in the collection of Mrs. E. G. Henriquez. The type locality
is from sands dredged near Habana and deposited at a Chorrera.

Range. Known only from the type locality.

Remarks. This is a very distinctive species and readily distinguished from all others
known in the Western Atlantic. It belongs to the group of D. aguayoi and D. wet-
moret, but is nearer to aguayoi. It differs from that species by having a smaller, broadly
oval rather than oblong orifice and is lacking the rounded tooth on either side of the ori-
fice. In addition, the shell of fragilis is more depressed, the radial ribs lower and more
widely spaced than in aguayoi.

Plate 179. Diodora fragilis Farfante and Henriquez. Holotype. Arenas de la Chorrera, Habana, Cuba.
Figs. 1-3 (23x). Fig. 4 (10x).

REFERENCES

Gray, J. E. 1854, List of the Shells of Cuba in the Collection of the British Museum, pp. 1-48.

Iredale, T. 1915, A Commentary on Suter’s Manual of the New Zealand Mollusea.* Transactions of the New
Zealand Institute 47, pp. 417-497.

Odhbner, N. 1932, Zur Morphologie und Systematik der Fissurelliden. Jenaische Zeitschrift fiir Naturwissen-
schaft 67, pp. 292-309, figs. 1-41.

344 JOHNSONIA, No. 39 Tricolia

Dene ee eee EE ee

Peile, A. J. 1926, Mollusca of Bermuda. Proceedings Malacological Society London 17, pp. 71-98.
Pilsbry, H. A. 1891, Manual of Conchology (1) 12, pp. 273-287.

Robertson, R. 1958, The Family Phasianellidae in the Western Atlantic. /ohnsoma 3, pp. 245-283, pls.
136-148.
Wenz, W. 1938 [in] Schindewolf. Handbuch der Paliozoologie 6, (Tp. £78:

Woodring, W. P. 1928, Miocene Mollusks from Bowden, Jamaica, Part II Gastropods. Carnegie Institution
of Washington, Publication no. 385, pp. 1-564, 40 plates.

* Kk Ck

THE GENERA AMAEA AND EPITONIUM IN THE
WESTERN ATLANTIC
BY
Rutu D. Turner

Amaea (Amaea) mitchelli

Amaea (Amaea) mitchelli Dall. Clench and Turner 1950, Johnsonia 2, p. 243, pl. 106, figs. 5-7.

Specimens examined. Texas: Mustang Island, 1 mile S of Port Aransas (Doil Turner).

Epitonium (Asperiscala) sericifilum Dall

Epitonium (Asperiscala) sericifilum Dall. Clench and Turner 1952, Johnsonia 2, p. 317, pl. 152.

Mrs. G. R. Hettick has kindly donated a specimen of this rare species to the Museum
of Comparative Zoology. Dall (1889, p. 124) in his list of the Marine Mollusks of the
Southeastern Coast lists this species with a question as occurring on the Texas Coast.
The specimen collected by Mrs. Hettick is from 10 miles south of Port Aransas, Texas.
It has 10 whorls and measures 6.5 mm. in length and 1.9 mm. in width.

THE FAMILY PHASIANELLIDAE IN THE WESTERN ATLANTIC
BY
Rovert ROBERTSON

Some additional observations on Tvicolia made at the Lerner Marine Laboratory,
Bimini, Bahama Islands in June and July, 1958, are included in this supplement, as well
as some new records from Cuba. Previously there were no records of T'ricolia on the
south coast of Cuba west of Guantanamo. The specimens were sent by Dr. C. G. Aguayo,
Museo Poey, Universidad de la Habana and Dr. Cortez Hoskins, Jersey Production
Research Co., ‘Tulsa, Oklahoma.

Tricolia affinis affinis C. B. Adams
Tricolia affinis affinis C. B. Adams. Robertson 1958, Johnsonia 3, no. 37, p. 262.
Specimens examined. Cusa: Cayo Inés de Soto; Playa Manimani, both Pinar del
Rio; Vedado; Arenas de la Chorrera, both Habana; Gibara, Oriente (all Museo Poey):

W of Cayo Tablones, Archipiélago de los Canarreos, Habana; Cayo Perro, Cayos de
San Felipe (both C. Hoskins); Pta. de la Yana, both Pinar del Rio (Museo Poey).

Western Atlantic JOHNSONIA, No. 39 345

Tricolia adamsi Philippi
Tricolia adamsi Philippi. Robertson 1958, Johnsonia 3, no. 37, p. 268.

Specimens examined. Cuna: Vedado, Habana (Museo Poey).

Tricolia thalassicola Robertson

Tricolia thalassicola Robertson 1958, Johnsonia 3, no. 37, p. 271.

Remarks. The median longitudinal furrow is present at the anterior end of the under
side of the foot only. Hence the waves of progression are only partially ditaxic. There
are irregular yellowish olive-green spots, somewhat as on the shell, on the upper surface
of the foot. The left cervical lobe is pedunculate and digitate, with four digits in two
specimens and three in another. The lobe on the right side is entire.

This species lives on Manatee Grass (Cymodocea manatorum Ascherson) as well as on
Turtle Grass (Thalassia) at Bimini. It was seen to feed on fine filamentous and crustose
calcareous red algae.

Specimens examined. Cuna: Cayo Inés de Soto; Playa Manimani, both Pinar del
Rio; S coast Camagiiey Province (subfossil; all Museo Poey); Cayo Perro, Cayos de
San Felipe (C. Hoskins); Pta. de la Yana, both Pinar del Rio (Museo Poey).

Tricolia bella MW. Smith

Tricolia bella M. Smith. Robertson 1958, Johnsonia 3, no. 37, p. 274.

Remarks. 'There is no median longitudinal furrow on the under side of the foot in this
species, as was previously stated to be characteristic of the whole genus (p. 250) and the
waves of progression are monotaxic, not ditaxic (see also Remarks above on 7°. thalassi-
cola). The earlier observation (p. 249) that there is no cervical lobe on the right side was
based on asingle abnormal specimen. Normally there is an entire lobe, as in 7°. thalassi-
cola. The left lobe had three digits in the specimens examined. The feces are U-shaped,
as in 7’. pullus.

Four young specimens (the largest 1.8 mm. long) were collected from floating Sar-
gassum west of Bimini. This species usually lives in mats of filamentous green algae.
Sometimes it may be found in the red alga Bostrychia {=‘‘Amphibia’’] tenella (Vahl)
Agardh on mangrove roots.

Specimens examined. Cuna: Vedado, Habana (Museo Poey); Cayo Perro, Cayos de
San Felipe, Pinar del Rio (C. Hoskins).

Tricolia tessellata Potiez and Michaud

Tricolia tessellata Potiez and Michaud. Robertson 1958, Johnsonia 3, no. 37, p. 277.

Range. This species was previously believed not to occur on the coast of Cuba. It has,
however, been collected on the south coast in a subfossil state. A record of this species
at Grand Cayman (Abbott 1958, Academy of Natural Sciences of Philadelphia Mono-
graph 11, p. 32) is based on a specimen collected alive of the banded form of 7°. thalas-
sicola (see Plate 146, fig. 3, p. 272).

346 JOHNSONIA, No. 39 Distorsio

Specimens examined. Cuna: S coast Camagiiey Province (subfossil; Museo Poey).

ADDITIONS AND CORRECTIONS

Page 280. The Carpenter specimens of Tricolia compta (Gould) are now at the United States National
Museum, Washington, D.C.

Page 280. Phasianella concolor C. B. Adams is probably a synonym of Assiminea succinea (Pfeiffer) [ Paludina
succinea Pfeiffer 1840]. Paludestrina auberiana d’Orbigny is not an Assiminea as was believed by Dall, but a

Littoridina.

Page 280. A. Heilprin 1893 (The Bermuda Islands, Philadelphia, p. 175) reports ““Phasianella Kochi, Phil.”’
from Bermuda. He evidently had some other species of Tricolia, for T. kochii (Philippi) is restricted to South
Africa (and possibly parts of the Indo-Pacific). There are three separate records of Tricolia from Bermuda
published during the last half of the nineteenth century and there is only one specimen of T. bella in the
Haycock Collection, Bermuda Government Museum (teste Richard W. Foster). None of the species appear to
have been collected in recent years. Evidently the genus is sporadic there.

Notes

Since the publication of Distorsio in this volume (pp. 235-242) we have obtained two
additional records of note.

Distorsio (Rhysema) clathrata Lamarck

Specimen examined. NortH Carouina: Combat, station 381, about 15 miles SE of
Cape Hatteras (N. Lat. 34°59’; W. Long. 75°33’) in 45 fathoms.

Distorsio (Rhysema) mcgintyi Emerson and Puffer

Specimen examined. BERMUDA: } mile S of Castle Rock in 80-100 fathoms (Bermuda
Biological Station).
This record extends the range considerably to the north.

Western Atlantic

JOHNSONIA, No. 39

347

INDEX
BY
Merritt E. CHampion

See T'able of Contents for references to new genera and species,
book reviews, voyages, and contributors.

The first page number given is to the most important reference.

abnormis, Scutum, 14

aborta, Pholadidea, 90

abyssorum, Xylophaga, 156,146

acclivis, Triton, 228

acinaces, Patella, 14

Acmaea, 14

acuminata, Parapholas, 128,9,123,
132

acuminata, Pholas, 128

adamsi, Phasianella, 268,274

adamsi, Tricolia, 268,263,270,
272,345

adamsii, Littorina, 262,274,275

affinis, Phasianella, 262,263

affinis, Tricolia, 262,248,251,253,
265,269,270,271,272, 344

aguayoi, Diodora, 343

aguayoi, Murex, 333

Aizyella, 252,261

alta, Pinna, 315,317,318

altispira, Melongena, 176,181,182

Amaea, 344

americana, Martesia, 102,104,107

americanum, Triton, 228,229

Anchomasa, 22

anomalus, Triton, 243

antillarum, Melongena, 164

antillarum, Murex, 333

antillarum, Triton, 225

anus, Distortio, 236

anus, Distortrix, 235

anus, Murex, 235,236

anus, Persona, 235

Apophyses, 13

aquatilis, Triton, 216

Aquilus, 200

Aquillus, 200

arconatii, Emarginula, 335

aspinosa, Melongena, 172,174

assimilis, Pinna, 314

Assiminea, 346

Astraea, 79

atlantica, Tritonia, 194

atlantica, Xylophaga, 152,5,146,
148,156

atlantis, Columbarium, 330

atomus, Pholas, 104

Atrina, 310,292,297,301

atropurpurea, Pinna, 290

auberiana, Assiminea, 280

auberiana, Paludestrina, 346

australasiae, Monoplex, 227,228

australe, Buccinum, 255,256

australis, Emarginula, 340

australis, Phasianella, 255,245,
248,249

Aviculopinna, 297,298

bahamensis, Martesia, 111,112,
13

Bailya, 243

Bankia, 3,146

barbudensis, Sconsia, 329

Barnea, 19,2,10,15,16,17

Barnia, 19

beaniana, Pholas, 104

beauii, Murex, 333

beaui, Tricolia, 263,265,269

beauiana, Pholas, 104

belknapi, Melongena, 172,174

bella, Phasianella, 274

bella, Tricolia, 274,249,250,251,
262,263,280, 345

bermudezi, Columbarium, 330

bicarinata, Phasianella, 280

bicarinata, Tricolia, 250

bicolor, Fusus, 182,184

bicolor, Melongena, 182,163,164,
168,178

bispinosa, Melongena, 180

bifrons, Pholas, 55,56

bispinosa, Pyrula, 180

bisuleata, Parapholas, 131,132

bitruneata, Panope, 50

Bolina, 255

branchiata, Parapholas, 182

brasilianum, Triton, 228,229

brayi, Columbarium, 330

brevis, Eucosmia, 258

brevis, Gabrielona, 258,248,251,
2575262

brevis, Phasianella, 257,258,262
263,268,270,272

brevis, Tricolia, 258

b]

brevissima, Phasianella, 258

Buccinatorium, 193

bulimoides, Phasianella, 256

Cabestana, 200

Cabestanimorpha, 200

Cadmusia, 89

caduceus, Murex, 200

Caducifer, 243

californianus, Mytilus, 79

californica, Parapholas, 124,70,130

californica, Pholas, 123,128

californica, Xylophaga, 154,155,
156

eallosa, Pholas, 45

eallosa, Zirfaea, 23

Callum, 13

ealva, Martesia, 131

calva, Parapholas, 131,123,128,
130

calva, Pholas, 131

cealypta, Tricolia, 253

campechiensis, Dactylina, 48

campechiensis, Pholas, 48,10,53,
170

campechiensis, Pragmopholas, 48

candeana, Pholas, 48,49

candida, Barnea, 19,10,20,24

eandida, Barnia, 20

eandida, Pholas, 19,21

earibaea, Pholas, 114,116

caribbaeum, Cymatium, 204,190,
208 ,242

earnea, Pinna, 306,286,287 ,288,
296,301

carolinensis, Pinna, 312

caudatus, Murex, 204

348

JOHNSONIA, No. 39

Index

Cerithium, 183,184

Chaceia, 66,158,3,16

Charonia, 193

Charonis, 193

chemnitzii, Triton, 243

chiloensis, Pholas, 51,10,49,50,
170

Chimaera, 301

Chimaeroderma, 301

chipolana, Tricolia, 253

chlorostomum, Cymatium, 190

chlorostomum, Triton, 210,211

Chromotis, 257,260

cingulata, Cassidaria, 242

cingulata, Linatella, 198

cingulatum, Cymatium, 200,242

clappi, Lignopholas, 98,99,100

clathrata, Distorsio, 236,235,240,
346

clathrata, Distortrix, 240

clathratum, Triton, 236

clathratus, Triton, 236

clausa, Emarginula, 336

clavata, Pholas, 101,102,103,107

clavator, Tritonium, 204

colosseus, Fusus, 187

Columbarium, 330

commutatus, Triton, 194

compta, Phasianella, 260,279,280

compta, Tricolia, 279,248,346

concamerata, Pholas, 80,83

concinna, Phasianella, 262,263

concinna, Tricolia, 268

concolor, Phasianella, 280,346

conoides, Pholas, 103

conradi, Penitella, 75,2,14,71,74,
84,122

constricta, Distorsio, 242

constricta, Zirfaea, 88

Conus, 329

corona, Melongena, 172,161,162,
163,164,170, 179,180, 182,184

corona, Murex, 172

coronatus, Fusus, 185

corrugatum, Cymatium, 221

corticaria, Pholas, 104,107

costata, Barnea, 9

costata, Cyrtopleura, 35,2,8,9,14,
40,42

costata, Holopholas, 35

costata, Pholas, 35

costata, Scobina, 35

costatus, Murex, 228,229,230

costatus, Pholas, 35

costulata, Pholas, 20

crassa, Phasianella, 261

crenulatus, Pholas, 23

crispa, Pholas, 55

crispata, Mya, 55

crispata, Pholas, 54,55

crispata, Zirfaea, 55,1,3,4,5,8,9,
10,11,21,24,30,59,61,91

crispus, Solen, 55

crucifera, Pholas, 34,41

cruciger, Cyrtopleura, 41,36,37,40

eruciger, Pholas, 34,41,42

crucigera, Pholas, 41

cruenta, Tricolia, 267,263,266,269

cumingii, Jouannetia, 9,134,135,
136

cuneiformis, Martesia, 114,6,50,
101

cuneiformis, Pholas, 114

cupula, Pholas, 104

Currus, 197

curta, Diplothyra, 121

curta, Martesia, 104,108

curta, Pholas, 121

curvata, Penitella, 80,83,84

curvata, Pholas, 81

cutaceum, Cymatium, 203

cutaceus, Aquillus, 200

cutaceus, Fusus, 198

cutaceus, Murex, 200

cutaceus, Triton, 198

cyanocephalum, Cymatium, 204

cyclostoma, Tricolia, 253

cylindrica, Barnea, 20

cylindrica, Pholas, 20

Cymatiidae, 189

Cymatilesta, 200

Cymatium, 231,197

Cymatriton, 210

Cynocephalum, 242

cynocephalum, Cymatium, 242,
198,200,206

cynocephalum, Triton, 204,205,
242

Cyrtopinna, 301

Cyrtopleura, 34,41,158,16,17

dactilus, Pholas, 45

Dactylina, 44

dactylina, Pholas, 45

dactyloides, Pholas, 19,23

dactylus, Dactylina, 45

dactylus, Hypogaeoderma, 45

dactylus, Pholas, 45,2,8,9,10,24,

dactylus, Pragmopholas, 45

darwinii, Netastoma, 141

darwinii, Nettastomella, 142,145

darwinii, Pholadidea, 142

darwinii, Pholas, 141,142,143

decorata, Phasianella, 256

decurtata, Pholas, 45,47

decussata, Pholas, 103

degenera, Pinna, 306

delicata, Diodora, 342

delicatula, Phasianella, 256

denudata, Melongena, 165,168

depressa, Emarginula, 336

dilecta, Pholas, 51,52,53

Diodora, 342

Diploplax, 102

Diplothyra, 118,158,6,101,140

Distorsia, 235

Distorsio, 235,236

Distorsus, 235

Distorta, 235

Distortio, 235

Distortix, 235

Distortrix, 235

divisus, Tagelus, 179

Dolarium, 200

d’orbignyi, Pinna, 315,317,318,
319

dorsalis, Teredo, 146

dorsalis, Xylophaga, 146,6,10,
149,150,153

dubia, Eudora, 262

duboisi, Pholas, 23

duchassaingi, Jouannetia, 135

effusa, Litiopa, 216

elongata, Pinna, 304,305

emarginata, Emarginula, 340

emarginata, Hemitoma, 340

Eotricolia, 247,252,261

Epitonium, 344

estephomenos, Hemifusus, 182

estephomenos, Melongena, 182

Eucosmia, 257,259,260

Eudora, 260

Eulithidium, 247,257,260,261,279

Eutriton, 193

Eutritonium, 193

Eutropia, 254,255

exaratum, Triton, 200

exigua, Phasianella, 261

exilis, Cyrtopleura, 41,42

exquisita, Martesia, 111,112,113

Western Atlantic

JOHNSONIA, No. 39

549

faleata, Pholas, 103,114,116

fasciata, Melongena, 165

fasciata, Pugilina, 184,185

felipponei, Cymatium, 203,201,
202

felipponei, Lotorium, 203

femorale, Cymatium, 232,189,231

femorale, Murex, 197,231,232
ferruginea, Pinna, 304,305
ficus, Murex, 165

finlayi, Murex, 331

fitchi, Penitella, 71,70
flabellum, Pinna, 307
flammeus, Turbo, 261

floridana, Distorsio, 239,240,241,

242

fordiana, Tricolia, 249

fosteri, Conus, 329

fragilis, Diodora, 342

fragilis, Martesia, 111,101, 102,
108,109

fulgens, Haliotis, 79

funisicola, Martesia, 104,107

fusco-nigricans, Pyrula, 168

gabbi, Penitella, 85,61,70,71,78,
84

gabbi, Tricolia, 253

gabbi, Zirfaea, 3,8,58,61,85,87

Gabrielana, 257

Gabrielona, 257,246,247 ,250,253,

256,272
galea, Tonna, 190
Galeodes, 165
gemmatum, Cymatium, 222,189
gemmatus, Triton, 222
gibbosa, Pholas, 148
gibbosus, Triton, 200
gigantea, Ranella, 228
Gitocentrum, 48
Glabella, 259
globosa, Jouannetia, 9,136
globosa, Triomphalia, 136
globosa, Xilotoma, 148
globosa, Xylophaga, 148,151
globosa, Xylotomea, 148
globulosa, Jouannetia, 138
globulosa, Pholadopsis, 136
goodall, Pholadidea, 89
goodall, Pholas, 90
gracilior, Strombus, 170
gracilis, Pholas, 45,47
gracilis, Triton, 222
guadaloupensis, Emarginula, 336

Gutturnium, 224

hadra, Tricolia, 253,257

haemastoma, Thais, 187

Hatasia, 92,89

hawaiiensis, Martesia, 104,107

Hemifusus, 187

Hemitoma, 336

hians, Pholas, 45,47

Hiata, 102

Hiloa, 247,252,261

Holopholas, 19

hornbeckii, Pholas, 104,107

Hypogaea, 44

Hypogaeoderma, 44

imperialis, Murex, 333

incisa, Penitella, 104

incurva, Pinna, 301

incurvata, Melongena, 172

infelix, Hiata, 102,104,105,107,
108

inflata, Atrina, 300

inspinata, Melongena, 172,174

intercalata, Martesia, 75,78,104,
108,122

intermedius, Triton, 216,217

intusgranosa, Mesopholas, 102,
104,107

Iouannetia, 133

Isognomonidae, 298

janellii, Pholas, 124,128

japonica, Atrina, 291

johnstonei, Melongena, 178,179,
163,164

jolyi, Tricoliella, 262

Jouannetia, 133,134,158,13,15,
16,17,140,146

Jouannetiinae, 16,133,17,65

klenei, Cymatium, 202

kochi, Phasianella, 346

kochii, Tricolia, 346

krebsi, Pholas, 114

krebsii, Cymatium, 220,223

krebsii, Triton, 220

labiosa, Murex, 201

labiosum, Cymatium, 201,190,200

labiosus, Murex, 201
laevigatum, Laevicardium, 319
lamarckii, Spondylus, 75
lamellata, Pholas, 90,91
lamellosa, Barnea, 25,22,24
lamellosa, Pholas, 25
lamellosus, Pholas, 90
Lampusia, 214

lanceolata, Barnea, 39

lanceolata, Cyrtopleura, 39,36,37,
43

lanceolata, Pholas, 39,40

laqueata, Pholas, 51,53

larvata, Penitella, 104

latior, Triton, 216

lehmanni, Phasianella, 256

ligamentina, Pholas, 23

lignivora, Penitella, 104

Lignopholas, 98,159,188

Linatella, 198

lineolatus, Phasianella, 256

lineolatus, Turbo, 256

lipeata, Phasianella, 277

listeri, Emarginula, 336

listeri, Pinna, 307,315,317

Littoridina, 346

longirostra, Ranularia, 204

loroisi, Cymatium, 202

loroisi, Triton, 200,201

loscombeana, Pholadidea, 90

loscombiana, Pholadidea, 90,89

Lotorium, 197

lotor, Lotorium, 197,232

lotorium, Triton, 232

lurida, Tricolia, 247

Luterium, 197

Mactesia, 101

Mactresia, 101

maculata, Macrocallista, 319

maculatus, Pinna, 296

major, Pholas, 23

margaritana, Melongena, 165,166,
168

margaritensis, Murex, 333

maritima, Barnea, 63,159

marmoratum, Tritonium, 193,196

Martesia, 101,102,158,1,2,15,16,
17,98,100,118,146

Martesiinae, 16,66,11,15,17,65

Martesiella, 102

martiniana, Melongena, 181

martiniana, Pyrula, 180

martinianum, Triton, 216

maura, Atrina, 291,318

346
melanura, Pholadidea, 93,9,92,96
melanura, Pholas, 92,93,94
Melongena, 165,163,164,170,175,
178
melongena, Galeodes, 165

$50

JOHNSONIA, No. 39

Index

melongena, Melongena, 165,167,
163,171

melongena, Murex, 165

melongena, Pyrula, 165

Melongenidae, 161

membranacea, Pinna, 298

Mesopholas, 105

Metaxylophaga, 145

mexicana, Xylophaga, 150,156

Mimelenchus, 255

minor, Ensis, 179

minor, Hemifusus, 182

minor, Melongena, 168,182

minuscula, Pholadidea, 111,113

minuta, Phasianella, 277

minuta, Tricolia, 278

minutissima, Eulithidium, 270

minutissimum, Eulithidium, 268

mitchelli, Amaea, 344

mohorteri, Cymatium, 243

mollis, Phasianella, 253

mollis, Tricolia, 253

Monoplex, 227

Monothyra, 44

Montfortia, 340

morio, Fusus, 185

morio, Melongena, 185

morio, Murex, 184,185

morio, Pyrula, 185

morio, Pugilina, 185,163

morio, Semifusus, 185

moritinetus, Cymatium, 208

moritinectus, Triton, 206

mulletensis, Melongena, 173

multispinosa, Melongena, 165,168

mundum, Triton, 222

Murex, 331

muricata, Pinna, 300,301,302,303,
320,321

muricatum, Trachycardium, 319

muricatus, Pholas, 45,47

muricina, Distorsio, 224,225

muricinum, Cymatium, 225,224

mutica, Pyrula, 187

Mytilidae, 298

Mytilus, 10

nana, Pholas, 131

nanus, Pholas, 103

Navea, 158

Neoxylophaga, 145

nepeanensis, Gabrielona, 258,246,
57

nepeanensis, Phasianella, 247,

25,208
Neptunella, 200
neritina, Chromotis, 260
neritina, Phasianella, 260
Netastoma, 141
Netastomella, 141
Nettastoma, 141
Nettastomella, 141,158,13,16,17,
134,146
newcombii, Navea, 75,77,78,79
nicobaricum, Cymatium, 210,190,
220,221
nicobaricum, Tritonium, 210
nigra, Pinna, 310
nobile, Triton, 193
nobilis, Pinna, 287,291,292,294,
301
nobilis, Triton, 194,196
nucicola, Mesopholas, 104,107
Nyctilochus, 193
obesa, Litiopa, 225,226
oblongata, Pholas, 48,49
oblongata, Thovana, 48
occidentale, Cymatium, 214,189
occidentale, Triton, 214
octoradiata, Emarginula, 340
octoradiata, Hemitoma, 336
octoradiata, Patella, 336
octoradiata, Subemarginula, 336
oldroydi, Atrina, 298
olearium, Monoplex, 227,228
olearius, Triton, 228
orientalis, Pholas, 14
orientalis, Triton, 201
Orthomesus, 255
Orthopnoea, 255
ostheimerae, Emarginula, 340
Ostreidae, 298
ostreum, Pinnotheres, 296
ovoidea, Chaceia, 66,74,84,87,88
ovoidea, Parapholas, 66
ovoidea, Pholadidea, 66,85,88
ovoidea, Pholas, 66,69
ovum, Pholas, 104
pacifica, Barnea, 31,33
Palaeopinna, 297,298
panza, Purpura, 170
papyraceus, Pholas, 90,91,19
paradisiaca, Volema, 165
Parapholas, 123,158,2,3,6,13,15,
16,17,148
Parlicymatium, 200
parthenopeum, Cymatium, 228,

189,227,230
parthenopeus, Murex, 228
Particoma, 114,102
Particymatium, 200
parva, Barnea, 23,22,33,55
parva, Barnia, 23
parva, Holopholas, 23
parva, Penitella, 14,75,77,78,79
parva, Pholas, 23,51,53
parvus, Pholas, 55
parvus, Triton, 243
patagonica, Pinna, 315
Patella, 14
patula, Cassidulus, 169
patula, Cyrtulus, 169
patula, Galeodea, 169
patula, Galeodes, 169
patula, Melongena, 168,166,167,

169,171
patula, Purpura, 170
patula, Pyrula, 168
paulucciae, Pinna, 304,305
pectinata, Jouannetia, 137,136,

140
pectinata, Pholadopsis, 136,137
Pellax, 246,247,252,261
Penicilla, 70
peninsulum, Cymatium, 198
penita, Penitella, 80,2,7,8,61,69,

70,71,74,78,79,87,88,128
penita, Pholadidea, 80
Penitella, 70,158,2,6,9
pennantiana, Anchomasa, 23
Pennaria, 310
pernula, Pinna, 304
Persoma, 235
Persona, 235
perspectiva, Melongena, 173
perspinosa, Melongena, 172,174
pharcida, Lampusia, 220
pharcidum, Cymatium, 220,221
Phasianella, 255,247,249,250,

252,254
phasianella ?, Trochus, 256
Phasianellidae, 245,254,344
Phasianellinae, 254
Phasianochilus, 261
Phasianus, 255
phasianus, Bulimus, 256
phasianus, Helix, 256
phaseolina, Penitella, 104
Pholadidae, 1,2,15,65
Pholadidea, 89,158,3,16,17,70,71

Western Atlantic

JOHNSONIA, No. 39

Pholadidoidea, 89

pholadiformis, Petricola, 50

Pholadinae, 15,1,11,19

Pholadopsis, 136,134

Pholas, 44,158,2,6,7,16,17,35

Phragmopholas, 44

picta, Phasianella, 256

pictus, Turbo, 261

pileare, Cymatium, 216,190,211,
215,220,221

pileare, Murex, 216

pileare, Triton, 216

pilsbryi, Zirfaea, 58,3,8,56,69

Pinna, 301,292,294,297,298

Pinnaria, 310

Pinnarius, 301

Pinnidae, 285,295,298

Pinnogena, 297

Pinnula, 301

pisum, Penitella, 104

Pododesmus, 79

pomum, Murex, 333

poulsenii, Cymatium, 198,190

poulsenii, Triton, 198

praestans, Xylophaga, 10

Pragmopholas, 44

precursor, Lacuna, 253

preissii, Phasianella, 256

prima, Dissentoma, 216,217

Prisogaster, 251

probrevis, Tricolia, 253,259

producta, Phasianella, 280

productum, Triton, 225

Protoplax, 12

Protoxylophaga, 145

Pseudophasianus, 252

Pteria, 298

Pteriidae, 298

pterocladica, Tricolia, 264,263,
266,268,269,278

Pugilina, 184,185,163,164

pugilis, Strombus, 170

pulchella, Littorina, 274

pulchella, Martesia, 104

pulchella, Phasianella, 256,261,
280

pulchella, Tricolia, 274

pulchella, Turbo, 270

pulcher, Murex, 333

pulchellus, Triton, 210,211

pulchellus, Turbo ?, 263,274,275

pulcherrima, Triomphalia, 136,
137,138

pullulus, Phasianella, 261
pullus, Phasianella, 261

pullus, Tricolia, 261,246,248,249,

250,252, 253,255,260, 345

pullus, Turbo, 260,261

pumilio, Triton, 210

punctata, Lacuna, 253

punctata, Tricolia, 253,261

pusillus, Pholas, 103,107

pygmaea, Phasianella, 267

pygmaea, Tricolia, 267,268

pyriformis, Triton, 225

pyruloides, Fusus, 187

pyrum, Cymatium, 209,210

pyrum, Murex, 225

quadra, Pholadidea, 95,92,94,97,
98

quadra, Pholas, 95

quadrangula, Barnea, 23

quillingi, Jouannetia, 139,134,
159,188

ramulosa, Pinna, 315,317

Ranula, 204

Ranularia, 204

rehderi, Cymatium, 208,206,211

reticulata, Distortrix, 236,239

retifer, Pholas, 51,53

retiporosa, Hemitoma, 340

retiporosa, Subemarginula, 340

Rexmela, 170

Rhysema, 236

ridleyi, Cymatium, 208

ridleyi, Triton, 206

rigida, Atrina, 312,287,289,290,
291,295,296,299,300,301,302,
311,318

rigida, Pinna, 312

rikuzenica, Xylophaga, 145

rivicola, Lignopholas, 98,100

rivicola, Martesia, 10

rosea, Pholas, 104,107

rostrata, Nettastomella, 14
141,142

rostrata, Pholadidea, 144,145

rostrata, Pholas, 143

rotunda, Distorta, 235

©9
to
fs

rubeculus, Murex, 214

rubens, Phasianella, 249,250,251,

255,256
rubida, Hemitoma, 336
rude, Chimeroderma, 304
rudis, Pinna, 304,293,295,300,
301,302,303,308

rufescens, Haliotis, 76,77,79

rugosa, Pinna, 291

rutilum, Tritonium, 201

sagitta, Pholadidea, 80,83,84

sarcostoma, Triton, 206

sarcostomum, Cymatium, 206,208,
210

sargenti, Melongena, 172

sarissophorum, Columbarium, 330

searlatina, Septa, 214

sciera, Hemitoma, 335

Scobina, 35

Scobinopholas, 35

Sconsia, 329

sculpturata, Melongena, 164

Scutum, 14

seebachi, Pinnogena, 297

semicaudata, Jouannetia, 133,134,
135

semicostata, Pholas, 159

semicostata, Scyphomya, 159

semicostata, Zirfaea, 159

Semifusus, 187

semispinosa, Melongena, 165,168

seguenzae, Tritonium, 194

seminuda, Atrina, 315,286,300,
301,302,303,305,307,309,311,
312,314,321, 322

seminuda, Pinna, 320,321

Septa, 214,193

sericiflum, Epitonium, 344

serrata, Atrina, 320,300,301,318

serrata, Pinna, 320

Servatrina, 314

setosa, Cassidaria, 243

shreevei, Capulus, 14,35,37

silicula, Penitella, 104

simillimus, Distorsio, 235

Simplum, 214

Simpulum, 214

Siphonella, 335

smithii, Diplothyra, 118,2,3,116,
123

smithii, Martesia, 118

solanderia, Cadmusia, 89,90

solanderiana, Pholadidea, 90

sorenseni, Haliotis, 79

spathulata, Barnea, 31,33

spathulata, Pholas, 31

spelaea, Penitella, 80

spengleri, Triton, 200

sphaeroidalis, Jouannetia, 140

spinosa, Barnea, 19

352

JOHNSONIA, No. 39

Index

squamifera, Atrina, 318

squamosissima, Pinna, 320,321

Steganomphalus, 260

stillata, Penitella, 104

strangei, Triton, 200,201,202

Streptopinna, 298

striata, Martesia, 103,1,3,6,8,65,
101,102,104,113,117,123

striata, Pholas, 90,102,103,106

striata, Sconsia, 329

subeoronata, Melongena, 170,172,
174,175

Subemarginula, 335

subglobosa, Navea, 75,78,79,158

subovata, Pholas, 20

subsanguinea, Phasianella, 256

subtruncata, Barnea, 31,22,27,29,

63,170

subtruneata, Pholas, 31

subviridis, Pinna, 309,315,317

succinctum, Triton, 228

succinea, Assiminea, 346

succinea, Paludina, 346

Sulcatopinna, 297

supplicata, Metaxylophaga, 145

syntoma, Tricolia, 253,257

Talona, 158,17

Talonella, 89

tenuis, Phasianella, 261

tenuistriata, Pholas, 103

Teredinidae, 1

teredinaeformis, Pholas, 104,107

terediniformis, Pholas, 104

Teredo, 3,146

tesselata, Phasianella, 263,277

tessellata, Phasianella, 277,280

tessellata, Tricolia, 277,262,263,
265,268,272,275,280,345

Thais, 187

thalassicola, Tricolia, 271,252,253,
262,263, 266, 278, 345

Thovana, 48,44,50

Thurlosia, 54

tigrinum, Cymatium, 233

tigrinus, Triton, 233,234

tokyoensis, Martesia, 104

tomlini, Xylophaga, 145

triangularis, Septa, 232

Tricola, 260

Tricolea, 260

Tricolia, 260,246,247 ,249,250,
252,253,255,256,257 , 344

Tricoliella, 260

Tricoliinae, 250

tricostata, Emarginula, 336

tricostata, Hemitoma, 336

tridens, Pholadidea, 159,91

tridens, Pholas, 159

tridens, Talona, 89,159

trinodulosa, Melongena, 173

Triomphala, 136

Triomphalia, 136

Triton, 189,193

Tritonalia, 193

Tritonellium, 193

Tritonia, 193

Tritonicus, 200

Tritonidea, 243

tritonis, Charonia, 195,196

tritonis, Murex, 193,196

tritonis, Phasianella, 256

tritonis, Tritonium, 193

Tritonium, 193

Tritonocauda, 204

Trochidae, 246

truncata, Barnea, 27,3,8,10,14,
22,24, 26,29,33,50,170

truncata, Pholas, 27,28

truncata, Zirfaea, 55,56,57

tuberculata, Pholas, 23

tuberculosa, Atrina, 314

tuberosum, Triton, 224,225

tubifer, Pholas, 96

tubifera, Pholadidea, 96,92

tubifera, Pholas, 96,97

Turbinidae, 246

turbinoides, Phasianella, 255

Turritriton, 200

typica, Phasianella, 279

typicum, Eulithidium, 279

umbilicata, Littorina, 262,271

umbilicata, Phasianella, 262,263,

271,275,280
umbilicata, Tricolia, 253
undosum, Triton, 243
Usatricolia, 260,261,279,280
varia, Phasianella, 256
varians, Eudora, 260,261 a
varicosa, Pinna, 295,304,305
variegata, Charonia, 193,190
variegata, Eucosmia, 260,279
variegata, Phasianella, 249,255,
256,279
variegata, Tricolia, 279,248
variegatum, Triton, 193,194
variegatus var. B, Triton, 194
veliei, Triton, 216,217
Venus, 10
venusta, Phasianella, 256
verrucosa, Hypogaea, 45
vexillum, Atrina, 290
vibonensis, Pholas, 90
vignoni, Jouannetia, 134,138,140
virginica, Crassostrea, 3,296
virgo, Pellax, 258
Volema, 165
waltonia, Didianema, 253,257
washingtona, Xylophaga, 154,153
waterhousii, Cymatium, 202
wetmorei, Diodora, 343
wiegmanni, Fusus, 242
wilsoni, Penitella, 93,94
xilophaga, Penitella, 104,107
xilophaga, Pholas, 146
Xilotoma, 145
Xilophaga, 145,158,1,2,9,10,15,
16,17
xylophaga, Penitella, 104
xylophaga, Pholas, 104
Xylophaginae, 17,145,11,15,65
Xylotomea, 145
Xylotrya, 44
zebrina, Phasianella, 262,277
Zinatella, 198
Zirfaea, 54,2,3,5,6,9,10,12,15,
16,17,57,68,70,88

BOUND

~

-

1975

(QQ

093 376 8

Date Due

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